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HARVARD UNIVERSITY 




LIBRARY 



OF THE 



Museum of Comparative Zoology 



,, ^^e Library 

••useu™ of Ccnaratlve Zoology 
Harvard University 



'- - i,iU;^. CU.ViP. ZCCL 

LIBRARY 

UNIVERSITY OF KANSAS miscellaneous 

MUSEUM OF NATURAL HISTORY JiW-il^Ajgpjj 

No. 55 

HARVARD^ 
UNlVERSl-ni 



Middle American Lizards of 
the Genus Ameiva (Teiidae) with 
Emphasis on Geographic Variation 



By 

Arthur C. Echternacht 



UNIVERSITY OF KANSAS 

LAWRENCE 1971 December 14, 1971 



Jk.. 



UNIVERSITY OF KANSAS PUBLICATIONS 
MUSEUM OF NATURAL HISTORY 

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Publications Secretary, Museum of Natural History, University of Kansas, Law- 
rence, Kansas 66044. 



University of Kansas 
Museum of Natural History 



Miscellaneous Publication No. 55 
December 14, 1971 



Middle American Lizards 

of the Genus Ameiva (Teiidae) 

with Emphasis on Geographic Variation 



By 

Arthur C. Echternacht 

Department of Biology 

Boston University 

Boston, Massachusetts 02215 



University of Kansas 

Lawrence 

1971 



University of Kansas Publications, Museum of Natural History 



Editors: Craig C. Black, William E. Duellman, 
Philip S. Humphrey 



Miscellaneous Publication No. 55 

pp. 1-86; 28 figures 

Published December 14, 1971 



Museum of Natural History 

University of Kansas 

Lawrence, Kansas 66044 

U.S.A. 



Printed by 

University of Kansas Printing Service 

Lawrence, Kansas 



CONTENTS 

INTRODUCTION 4 

ACKNOWLEDGMENTS 5 

MATERIALS AND METHODS 6 

Characters 6 

Statistical Procedures 11 

Sexual Dimorphism 1 2 

Sources of Specimens Examined 12 

SYSTEMATIC ACCOUNT OF THE SPECIES 13 

KEY TO THE MIDDLE AMERICAN SPECIES OF AMEIVA 13 

Ameiva ameiva ( Linnaeus ) 14 

A meiva leptoph njs Cope 20 

Ameiva f estiva ( Lichtenstein and Von Martens) 26 

Ameiva quadrilineata (Hallowell) 34 

Ameiva undulata ( Wiegmann ) 40 

Ameiva chaitzami Stuart 59 

DISCUSSION ■ 63 

INTERSPECIFIC RELATIONSHIPS 63 

INTERSPECIFIC PATTERNS IN GEOGRAPHIC VARIATION 64 

HISTORY OF AMEIVA IN MIDDLE AMERICA 67 

SUMMARY 71 

RESUMEN 72 

APPENDIX A: SUMMARY OF STATISTICS 75 

APPENDIX B: SPECIMENS EXAMINED 76 

LITERATURE CITED 83 



INTRODUCTION 



In their revision of the Mexican 
species of Ameiva, Smith and Laufe 
( 1946 ) prefaced their comments with 
the remark that interest in these species 
had "been rising to a crescendo whose 
peak surely could soon be anticipated." 
Little could they have foreseen the 
surge of interest, both taxonomic and 
ecological, that the entire genus has 
enjoyed over the past five years. This 
attention has been precipitated by the 
realization that Arneiva, by virtue of its 
wide distribution in the Neotropical 
Realm and the extremes of variation 
exhibited by its species, is especially 
well suited for ecological, biogeograph- 
ical, and evolutionary studies. Coupled 
with this has been the discovery that 
some species of Ameiva act as vectors 
for disease-causing organisms actually 
or potentially dangerous to humans (for 
example, see Schneider, 1965). 

There are three natural geographic 
groupings of Ameiva: those found on 
the various islands of the Caribbean, 
those on the continent of South Amer- 
ica, and those from Middle America 
(Mexico and Central America). Taxo- 
nomic studies of the Caribbean Arneiva 
have been conducted by Baskin and 
Williams ( 1966 ) , Heatwole and Torres 
(1967), Schwartz (1965, 1967, 1968), 
Schwartz and Klinikowski ( 1966 ) , and 
Schwartz and McCoy (1970). Very 
little recent work has been done on the 
Ameiva of South America; the only 
study is that of Peters (1964) on the 
Ecuadorian species. No recent compre- 
hensive taxonomic studies have been 
made of the Middle American species, 
although Stuart (1942) and Smith and 
Laufe ( 1946 ) contributed to our knowl- 
edge of certain species. The most re- 
cent revision of the entire genus is that 
of Barbour and Noble (1915). Their 
conclusions concerning the Middle 
American species of Ameiva were based 
on examination of only 41 specimens, a 
situation which largely masked the re- 



markable degree of geographic varia- 
tion exhibited by some of the species. 
It is the purpose of this study to present 
an analysis of the geographic variation 
in each of the Middle American species 
of Ameiva as well as to summarize 
known information on the biology and 
ecology of these lizards. 

Some persons will look upon the 
present work as justification for the rec- 
ognition of subspecies of Middle Amer- 
ican Ameiva. The controversy over sub- 
species has been debated extensively 
and heatedly elsewhere (Wilson and 
Brown, 1953; Peters, et ah, 1954; Smith 
and White, 1956; Savage and Heyer, 
1967; Smith, 1967; and others), and I 
will not dwell on it here other than to 
state my conviction that a thorough de- 
scription of geographic variation may 
make unwarranted and often undesir- 
able the arbitraiy nomenclatural frag- 
mentation of highly variable species. 

There is a growing body of evi- 
dence indicating the congeneric status 
of Ameiva and Cnemidophorus. Van- 
zolini and Valencia ( 1965 ) , Estes 
(1969), and Gorman (1970) demon- 
strated the closeness of the relationship 
and discussed the difficulties encoun- 
tered in trying to diagnose the two 
genera on the basis of known charac- 
ters. Gorman (1970:240), however, was 
hesitant to make a decisive statement 
on the matter when he commented: 
"The similarity of karyotypes between 
Ameiva and the southernmost Cnemido- 
phorus species group does not neces- 
sarily demonstrate close relationship, 
but there seem to be no characters that 
clearly separate the two genera." I be- 
lieve that the argument will not be re- 
solved until increased knowledge of 
South American Ameiva, especially the 
perplexing A. lacertoides, is available. 
The condition of the base of the tongue 
(free and heart-shaped in Cnemido- 
phorus, sheathed in Ameiva) has been 
considered to be diagnostic (Burt, 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



1931), but A. lacertoides has a Cnemi- 
dophorus-\ike tongue (Vanzolini and 
Valencia, 1965; personal observation). 
I prefer not to comment on the issue, 
but instead await the completion of 
relevant investigations by Richard Estes, 
George C. Gorman, William P. Mac- 
Lean III, and William Presch. 

ACKNOWLEDGMENTS 

During the course of this study, I 
have built up a huge debt of gratitude 
to many people who have provided as- 
sistance in one way or another. I have 
probably taxed the patience of a num- 
ber of museum curators, curatorial as- 
sistants, and collectors with requests for 
the loan of large numbers of specimens 
and locality or ecological data on spe- 
cific specimens. I especially acknowl- 
edge the following: Kraig K. Adler, the 
late Doris M. Cochran, James R. Dixon, 
William E. Duellman, Henry S. Fitch, 
George Foley, Jean Guibe, Giinther 
Peters, Robert F. Inger, Daniel A. Jan- 
zen, John M. Legler, Charles H. Lowe, 
Edmond V. Malnate, Hymen Marx, 
John R. Meyer, Charles W. Myers, 
Thomas Olechowski, James A. Peters, 
Douglas C. Robinson, Richard D. Sage, 
Jay M. Savage, Norman J. Scott, Hobart 
M. Smith, Dorothy M. Smith, A. G. 
Stimson, L. C. Stuart, Stephen G. Til- 
ley, Donald W. Tinkle, Robert G. Tuck, 
Jaime Villa, Ernest E. Williams, John 
W. Wright and Richard G. Zweifel. 

I am indebted to Richard F. John- 
ston, Irving Pfau, James D. Rising, F. 
James Rohlf, Gerald R. Smith, and 
especially Dennis M. Power for advice 
on statistical methods or computer pro- 
cedures. The computer programs uti- 
lized in this study were provided by 
Dennis M. Power. 

I have had the benefit of a great 
deal of constructive criticism and advice 
from my colleagues at the University of 
Kansas Museum of Natural History. 
William E. Duellman has been a con- 
stant source of encouragement, and his 
patience with me has, at times, been 



remarkable. He, along with Henry S. 
Fitch and Ronald L. McGregor, have 
reviewed this manuscript. Donald R. 
Clark, Ko Ko Gyi, John D. Lynch, Rus- 
sell J. Hall, Charles W. Myers, E. H. 
Taylor and Linda Trueb have been will- 
ing to discuss special problems as they 
arose, and their help is gratefully 
acknowledged. 

Richard Estes, George C. Gorman 
and James A. Peters have kindly al- 
lowed me to read their unpublished 
manuscripts, and Estes has also been 
gracious with his time and knowledge 
of fossil teiids. 

E. Raymond Hall, Philip S. Hum- 
phrey, and William E. Duellman of the 
University of Kansas Museum of Nat- 
ural History have allowed me free ac- 
cess to the collections and facilities' of 
the museum. The Panamanian collec- 
tions made by Duellman and Charles 
W. Myers have been especially useful. 
Field work was financed in part by Na- 
tional Science Foundation Summer Fel- 
lowships for Graduate Teaching As- 
sistants ( 1965 and 1966 ) and by travel 
grants and traineeships provided by the 
Committee fof Systematics and Evo- 
lutionary Riology from funds granted 
by the National Science Foundation 
(Grant No. GR-4446X) to the Univer- 
sity of Kansas, Dr. William A. Clemens 
and Dr. George W. Byers, principal in- 
vestigators (1966-1968). A grant-in-aid 
from the Society of the Sigma Xi pro- 
vided funds for illustrations, which were 
executed by Mrs. Susann Jacobshagen. 
Computer-time was made available by 
the University of Kansas and Boston 
University Computer Centers. 

To my wife Sandra, I owe my great- 
est thanks. She has been a constant 
source of encouragement and support 
and has proven to be an able assistant 
in both field and laboratory. She has 
cheerfully endured the many incon- 
veniences experienced by the families of 
graduate students, and, with the realiza- 
tion that the debt can never be repaid, 
I dedicate this work to her. 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



MATERIALS AND METHODS 

This study is based on the examina- 
tion of over 3000 specimens of Ameiva, 
2848 of which were examined in detail 
with respect to morphology, color and 
pattern. Samples were drawn from 
throughout the ranges of Ameiva fes- 
tiva, A. leptophrys, A. quadriUneata, A. 
tindulata, and A. chaitzami. Onlv Pan- 
amanian specimens of Ameiva ameiva 
were examined, because to include 
specimens from throughout the range 
of the species would have necessitated 
examination of material from South 
America, where inadequate geographic 
representation of samples would have 
made any conclusions more than usually 
speculative. Ameiva ameiva will be in- 
cluded in a projected study of South 
American members of the genus which 
will be completed only after field work 
necessary for the accumulation of sam- 
ples representative of the entire South 
American range of the species is pos- 
sible. South American samples of 
Ameiva festiva have been included in 
the present study because the range of 
the species is primarily in Central 
America. 

For purposes of statistical analysis, I 
attempted to amass samples of at least 
30 specimens (ideally 15 males and 15 
females) from relatively restricted 
localities. This attempt was only par- 
tially successful in that, with the excep- 
tions of Guatemala and parts of Mexico, 
large series of Ameiva from single lo- 
calities are not available. Of necessity, 
samples of ten or more have been used; 
for characters showing sexual dimor- 
phism (Table 1), the samples were still 
smaller. Greater success was achieved 
in accumulation of samples representa- 
tive of the ranges of the species, al- 
though lack of collections from critical 
areas in Panama has precluded exact 
statements of variation in A. ameiva 
and, especially, A. leptophrys. 

In each of the species accounts, a 
short synonymy, statement of distribu- 
tion, diagnosis, and description are fol- 



lowed by an extensive discussion of 
infraspecific geograpliic variation, eco- 
logical notes, and general remarks. A 
discussion of interspecific relationships 
and the history of the genus Ameiva in 
Middle America follows the species 
accounts. 

Characters 

In the interests of standardization of 
methods, a definition of each of the 
characters analyzed in this study fol- 
lows. Unless stated otherwise, charac- 
ters are defined in the same manner as 
by Smith ( 1946 ) . An asterisk ( * ) indi- 
cates a character analyzed for geo- 
graphic variation by Power's (1970) 
modification of Gabriel's ( 1964 ) Sums 
of Squares Simultaneous Test Proce- 
dure (STP). Interpretation of the re- 
sults of STP analysis is summarized in 
the section on statistical procedures. It 
will be noted that certain characters 
were not taken for some species, where- 
as in other instances a character was 
taken but not analyzed by the STP 
method. 

Snoiit-Vent Length (SVL). — The 
distance, to the nearest millimeter, from 
the tip of the snout to the anterior mar- 
gin of the vent. Maximum observed 
values for males and females of each 
species are given. Collections seem to 
be biased in favor of larger (male) in- 
dividuals. Measurements were taken 
with a flexible plastic rule. 

Head Length. — The distance to the 
nearest 0.1 millimeter from the tip of 
the snout to the posterior surface of the 
quadrate bone. The latter point was 
found by piercing the tympanum with 
one tip of the dial calipers and hooking 
the tip behind the bone. That the quad- 
rate was, in fact, the bone encountered 
was verified by x-ray photography. 

^Supralahials. — The total number of 
supralabials on both sides of the head. 
The posterior supralabial is herein de- 
fined as the last enUu'ged scale in con- 
tact with both the lip and a subocular 
scale (Fig. la). 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



'^Infralabiah. — The total number of 
infralabials on both sides of the head. 
The posterior infralabial is determined 
by the point at which the masseter 
muscle and associated membranes pass 
from the lower to the upper jaw. 

'* Terminal Siiblabials. — This charac- 
ter was noted for Ameiva festiva only. 
Stuart (1943:21) utilized the number 
of terminal sublabials to distinguish A. 
festiva festiva from A. /. edwarchi. I 
counted the sublabials present in the 
position of the two terminal sublabials 
of a "typical" A. /. festiva, totaling the 
numbers obtained for both sides of the 
head. By this method, a "typical" A. /. 
festiva (sensu Stuart) has four, whereas 
A. /. echvardsi has six or more. 

'* Supraoculars. — The total number 
of supraoculars on both sides of the 
head. Supraoculars are numbered from 
front to back (3a-c, Fig. lb); there are 
three or four on each side of most 
specimens of Ameiva from Middle 
America. 

'^Loreals. — The total number of 
loreals on both sides of the head. If 
more than one loreal is present on one 
side of the head, the second is smaller 
and usually located at the angle formed 
by the contact between supralabials 
and suborbitals anterior to the eye (3, 
Fig. la). Occasionally, there are two 
loreals about equal in size, one dorsal to 
the other. If a third scale is present in 
the area encompassed by the loreal(s), 
it is counted as a third loreal only if it 
is one-half the size of the smaller of the 
two other loreals or larger. 

^Degree of Circumorbital Granule 
Contact With the Frontal (COF).— The 
degree of contact by the circumorbital 
granular scales with the frontal was 
coded for purposes of statistical analysis 
as follows: 1) circumorbital granules 
not in contact with frontal-frontopari- 
etal suture (Fig. lb), 2) circumorbital 
granules in contact with frontal-fronto- 
parietal suture, but not extending past 
it (Fig. Ic), 3) circumorbital granules 
extending forward past the frontal- 



frontoparietal suture (Fig. Id). The 
coded values obtained for both sides of 
the head were totaled. Thus, the low- 
est possible value for a given specimen 
is two; the highest is six. 

"^Circumorbital Pattern (COP). — The 
position of the anteriormost circum- 
orbital granule relative to the supra- 
ocular scales was coded as follows: 1) 
granules extending to the level of the 
middle of the first supraocular scale, 
2) granules extending to the level of 
the suture between the first and second 
supraocular scales, 3) granules reach- 
ing the level of the middle of the sec- 
ond supraocular scale (Fig. Id), 4) 
granules extending to the level of the 
suture between the second and third 
supraocular scales (Fig. Ic), 5) gran- 
ules extending to the level of the mid- 
dle of the third supraocular scale (Fig. 
lb), 6) granules extending to the level 
of the suture between the third and 
fourth supraocular scales or, if there is 
no fourth supraocular, granules not 
reaching the level of the middle of the 
third supraocular, 7) granules extend- 
ing to the middle of the fourth supra- 
ocular scale. Coded values for both 
sides of the head were totaled for 
analysis. 

'^Extent of the Double Row of Gran- 
ules Between the Supraocular and Su- 
perciliary Scales (SO-SC). — This char- 
acter was coded as follows: 1) single 
row of granules between the super- 
ciliary and supraocular series ( Fig. Id ) , 
2) double row of granules extending 
foi-ward to the level of the middle of 
the third supraocular scale, 3) double 
row of granules extending forward to 
the suture between the second and third 
supraocular scales, 4) double row of 
granules extending forward to the 
level of the middle of the second supra- 
ocular (Fig. Ic), 5) double row of 
granules completely separating the su- 
perciliary scales from the second and 
third supraocular scales (Fig. lb). 
Coded values for both sides of the head 
were totaled for analysis. 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 







B 




Fig. I. Details of head and preanal scutellation of Ameiva. (A) Lateral view of 
head: 1. — Postnasal; 2. — Anterior loreal; 3. — Posterior loreal; 4. — Suborbital 
scale in contact with posterior supralabial (small, rectangular scale below 4). 
(B, C, and D) Dorsal view of head, left supraorbital region: 1. — Superciliary 
series; 2. — Supraocular-superciliary granules; 3A, 3B, and 3C. — Anterior, mid- 
dle and posterior supraocular scales, respectively; 4. — Prefrontal scale; 5. — 
Frontal scale; 6. — Frontoparietal sca'e; 7. — Circumorbital granules. (E) Pre- 



anal region: 1.- 



-Anterior 
scale; 3.- 



(1st) preanal 
—Terminal row 



scale; 2. — Posterior (last) preanal 
of ventral scutes. 



Gular Scales. — The central gular 
scales are small and irregular in ar- 
rangement, large and irregular, or large 
and oriented longitudinally in the mid- 
line. Intermediate conditions exist, but 
most specimens can be assigned without 



difRculty. If enlarged midgular scales 
are present, the smaller scales around 
them may become gradually smaller to- 
ward the periphery of the gular region 
or may become gradually smaller 
laterally and anteriorly but abruptly 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



9 



very small posteriorly. If the latter is 
true, the gular region is distinctly di- 
vided into pregular and postgular areas 
(see Fig. 5c, A. leptophrys) . 

MesoptychiaJ Scales. — The meso- 
pt\'ehial scales are small and irregular, 
continuous with the posterior gular 
scales, or they may form a band of 
abruptly enlarged scales across the 
throat. 

"^Granules Around the Body (GAB). 
— The number of granules around the 
body was shown by Zweif el ( 1959 ) to 
be of diagnostic importance in distin- 
guishing species of Cnemidophorus, and 
this character has been used extensively 
by students of that difficult genus. In 
most cases, the character has been given 
as "scales around midbody" (I believe 
the term "granules" to be more de- 
scriptive), but rarely is there a state- 
ment as to what "midbody" refers. This 
is important inasmuch as the character 
is intended to be a measure of granule 
size, and the granules vary in size over 
the length of the body. In this study, 
GAB counts were made starting at a 
point adjacent to the 15th transverse 
row of enlarged ventral scutes, the first 
transverse row of ventral scutes is taken 
as the first row extending entirely across 
the chest behind the gular fold. 

*" Paravertebral Granules (PV). — The 
number of granules between, and in- 
cluding, the paravertebral stripes of 
Ameiva leptophrys, quadriUneata, un- 
dulata, and chaitzami. 

'^Vertebral Stripe (VS). — The num- 
ber of granules included within the 
vertebral light stripe of Ameiva festiva. 

PV and VS counts were omitted for 
A. ameiva because the stripes are lack- 
ing or very faint on most adults of that 
species. 

""PV/GAB Ratio.— The ratio of para- 
vertebral granules, as defined above, to 
granules around the body was calcu- 
lated. 

^Granules Occiput to Rump (GOR). 
— The number of granular scales be- 
tween the enlarged occipital scales and 



the large, keeled dorsal caudal scales 
was counted for all species. Although 
there is usually a high correlation be- 
tween values obtained for this count 
and those for granules around the body 
(correlation coefficient=0.75 and 0.54 
for males and females of A. leptophrys, 
respectively), GOR sometimes shows 
sexual dimorphism and occasionally is 
an easier and more accurate count than 
GAB. The latter is especially true in 
poorly preserved specimens that have 
been coiled or that have numerous folds 
in the skin of the flanks. With experi- 
ence, neither GAB nor GOR counts are 
difficult or time-consuming to make, 
and error is not excessive (2-3% based 
on recounts of the same specimens). 
The most suitable technique for count- 
ing dorsal granules seems to be to use 
a fine insect pin as a pointer and marker 
while counting granules beneath a wide- 
field, binocular microscope. Ideally, the 
specimens should be slightly dry, since 
fluid between the granules results in 
glare and makes counting difficult. 

''GAB /GOR Ratio.— This ratio was 
calculated for all species. 

Prebrachials. — The number of rows 
of prebrachials was noted. 

Posthrachials. — T h e postbrachials 
are enlarged or not. 

Preantebrachials. — The number of 
rows of preantebrachial scales was re- 
corded. 

Prefe moral Scales. — Prefemoral scales 
are defined herein as the large scales 
on the anterior and ventral surfaces of 
the upper hind leg as far back as the 
femoral pore series. Thus defined, pre- 
femorals include the infrafemoral and 
prefemoral scales of Smith (1946:28). 
The number of rows was counted. 

Infratibial Scales. — The number of 
rows of infratibial scales was counted. 

The number of rows of preante- 
brachials, prebrachials, prefemorals and 
infratibials is variable throughout the 
length of the series on a single speci- 
men. In many specimens, there is a 
greater number of scales medially than 



10 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



proximally. This variation is discussed 
for each species. 

'* Femoral Pores. — The number of 
femoral pores was recorded; counts 
given in the text and tables are for the 
total number of pores on both legs. 

'^Subdigital Lamellae. — The number 
of subdigital lamellae on the anterior 
margin of the fourth (longest) toe of 
the left hind foot was recorded. The 
starting point for this count is the 
tubercule immediately proximal to the 
enlarged tubercule at the base of the 
first phalanx. If there is no tubercule 
in this position, the enlarged basal 
terminal is itself used as the starting 
point. 

'* Longitudinal Rows of Ventral 
Scutes. — The number of rows of en- 
larged ventral scutes was counted be- 
tween the gular fold and the first pre- 
anal scale, as defined below. The first 
row is the anteriormost row extending 
entirely across the chest posterior to 
the gular fold. 

'^Transverse Rows of Ventral Scutes. 
— The number of transverse rows of 
ventral scutes was counted at the level 
of the 15th longitudinal row of ventrals. 
The outer row on each side is reduced 
in size in some specimens. 

'^ Total Preanal Scales. — The number 
of preanal scales from just posterior to 
the tenninal row of longitudinal ven- 
tral scutes to the enlarged scale(s) pre- 
ceding the vent were counted. The 
first preanal scale is usually single, lies 
between the proximal femoral pores of 
each leg, and is distinguished from the 
last row of ventrals by its unpaired con- 
dition (1, Fig. le). Smith and Laufe 
(1946:20) encountered difficulty in de- 
termining which preanal scale is termi- 
nal because the preanal scales occa- 
sionally grade in size into the granular 
scales immediately anterior to the vent. 
When in doubt as to which preanal or 
row of preanals is terminal, I have sim- 
ply recorded the last row having three 
or fewer enlarged scales. In most cases, 
this is obviously the terminal row any- 



way. In those lizards with a single 
terminal preanal, the scales lateral to it 
are usually very much smaller and 
create no confusion as to whether or 
not they are part of the terminal pre- 
anal series. 

'^Termitml Preanal Scales. — The 
number of scales in the temiinal posi- 
tion in the preanal series was deter- 
mined by the method outlined above 
(2, Fig. le). 

''Scales Around the Tail (SAT).— 
The number of scales around the tail 
was counted at the level of the 15th 
verticil of caudal scales; the first verticil 
was taken as that row of caudal scales 
which first completely encircles the tail 
posterior to the vent. 

Various aspects of color and color 
pattern were noted on a sample-by-sam- 
ple basis. Recurring pattern compo- 
nents noted were as follows: 

Vertebral Stripe. — A narrow stripe 
originating on the snout or occiput and 
terminating on the tail unless com- 
pletely or partially lost in adults. 

Middorsal Field or Area. — A broad 
stripe, containing a vertebral stripe or 
not, originating on the occiput and 
terminating on the tail, bordered later- 
ally by the dorsolateral light stripes, if 
present, or the dorsolateral dark field. 

Dorsolateral Light Stripe. — A nar- 
row, light-colored stripe originating on, 
or posterior to, the superciliary rows 
and terminating on the tail and which 
may be entirely or partially lost in 
adults. 

Secondary Stripe. — A narrow, dark- 
colored stripe, the lateral border of 
which is the dorsolateral light stripe, if 
present, or the dorsolateral dark field 
and the medial border is the lighter part 
of the middorsal field in which it lies. 

Dorsolateral Dark Field. — A dark, 
often black or dark brown, area 
bounded above by the dorsolateral light 
stripe, if present, or the middorsal field 
and below by the ventrolateral light 
stripe, if present, or the ventrolateral 
field; originating on the loreal scales, 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



11 



terminating on the tail and interrupted 
by the eye; may be indistinct or lost in 
adults. 

Ventrolateral Light Stripe. — A nar- 
row, light-colored stripe originating at 
the posterior margin of the eye or pos- 
terodorsal margin of the ear and ex- 
tending along the body to the groin, 
usually continuing behind the leg and 
slightly onto the tail. 

Ventrolateral Field or Area. — Ven- 
trolateral one-half or two-thirds of the 
flanks bordered above by the ventro- 
lateral light stripe, if present, or the 
dorsolateral dark field and below by the 
enlarged ventral scutes; distinguished 
from the dorsolateral dark field by 
its generally lighter color; may contain 
irregularly arranged spots, blotches 
and/ or bars or regularly spaced vertical 
bars. 

Lateral Bars or Blotches. — Regular 
blotches in the dorsolateral dark field 
or bars on the flanks. 

Head scutellation is extremely vari- 
able in Ameiva. This is especially evi- 
dent in Ameiva leptophrys and, to a 
lesser extent, in festiva and quadriline- 
ata. The supraoculars of specimens of 
quadrilineata from the Bocas del Toro 
region of Panama are so fragmented as 
to preclude counting them. There may 
be three or four supraoculars per side 
in any of the Middle American species 
of Ameiva, and specimens of all species 
are known which have four on one side, 
three on the other. The number of 
supraoculars is thought to be sufficiently 
constant in Cnemidophorus that the 
number is used to distinguish species 
groups ( Burt, 1931 ) . The posterior dor- 
sal head scales of Ameiva leptophrys 
(Fig. 7) and some festiva are variously 
fragmented, so that in extreme cases 
parietal scales are not recognizable, and 
the frontoparietals and frontal are only 
slightly less disrupted. It might seem 
that such variability in the posterior 
head scales of Ameiva renders them 
useless for taxonomic purposes, but it 
seems that the amount of variation is a 



useful taxonomic character. For exam- 
ple, the presence of small, irregular 
scales separating the inteiparietal from 
the parietals and frontoparietals can be 
used to distinguish leptophrys from 
other Middle American Ameiva. 

Statistical Procedures 

Statistical summaries of all species 
examined comprise Appendix A. 

Power's ( 1970 ) modification of Gab- 
riel's (1964; see Gabriel and Sokal, 
1969) Sums of Squares Simultaneous 
Test Procedure ( STP ) provides a test 
of significance or non-significance of 
differences among sample means and is 
employed only after an overall analysis 
of variance demonstrates significant dif- 
ferences among the means. Samples are 
then ranked in decreasing order of their 
means and sums of squares calculated 
by sequentially adding means (be- 
ginning with the largest) until a maxi- 
mal non-significant subset is delimited. 
The procedure is then repeated, delet- 
ing one or more of the larger means in 
the previously defined subset, until 
another maximal non-significant subset 
is described. This procedure is repeated 
until all samples have been included in 
at least one such subset. A constant 
probability level of 0.05 was used in the 
study, as recommended by Power 
( 1970 ) . Calculations were performed 
by the University of Kansas Computa- 
tion Center on a GE-625 computer, and 
at Boston University on an IBM 360-40. 

In addition to STP, a number of 
statistics were calculated for each spe- 
cies and each sample. Certain of these, 
along with the STP results, are sum- 
marized in the tables accompanying 
each species account. Information per- 
taining to the total number of femoral 
pores of Panamanian Ameiva ameiva 
will serve as an example for the inter- 
pretation of STP results (see Appendix 
A). The first column of numbers in- 
cludes the locality code, keyed to the 
range map of Panamanian A. ameiva 
(Fig. 2). The localities are ordered in 



12 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



TABLE 1. — Occurrence of sexual dimorphism among Middle American Atneiva as determined 
by STP analysis. A, ameiva; L, Icptophnjs; Q, quadrilineata; F, festiva; U, undulata; C, chait- 

:ami: N, not tested. 



Species 

Character ~A L Q F U C~ Character 

Supralabials — + — + + — PV/GAB 

Infralabials ______ GOR 

Supraoculars ______ GAB/GOR 

COF values +_____ SAT 

COP \'alues ______ Total Preanals 

SO-SC values N — N N — — Terminal Preanals 

Loreals N — N N — — Subdigital Lamellae 

Sublabials N N N + N N Trans. Rows of Ventrals 

GAB _ _ _ _^ _ _ Long. Rows of Ventrals 

PV N — — + + — Femoral Pores 



Species 



Q 



u 



N 



- - + - 



+ 



+ 



+ 


— 


+ 


— 


+ 


— 


— 


— 


+ 


— 


+ 


— 


— 


+ 


+ 


+ 


+ 


— 


+ 


+ 


+ 


+ 


— 


— 


— 


— 


+ 


— 


— 


— 



+ + + + + 



terms of tlie decreasing magnitude of 
their means, given in column two. Fol- 
lowing the mean values are the standard 
deviation of the mean, sample size (in 
parentheses), and observed range. To 
the right of each block of statistics are 
the STP results, appearing as vertical 
lines. There are no STP results given if 
the overall analysis of variance indi- 
cated no significant differences among 
the sample means, and in these in- 
stances the samples are ordered by their 
locality numbers irrespective of their 
mean values. In the example, samples 
6 and 5, with means of 35.0 and 34.1, 
respectively, form the first maximal non- 
significant subset and are so indicated 
by the left-most line. Localities 5, 7, 3, 
2 and 1 constitute the second subset 
(middle line), and localities 7, 3, 2, 1 
and 4 are the third (right line). The 
mean for locality 6 is significantly dif- 
ferent from all other means, except that 
of locality 5 as indicated by the obser- 
vation that the mean for locality 6 ap- 
pears in no subsets containing means 
other than that of locality 5. Likewise, 
locality 5 is significantly different from 
locality 4. Localities 5 and 7 are not 
statistically different, although it would 
appear so if only the third subset were 
considered, but in this subset locality 5 
was omitted from the comparison in 



order to allow other means into the sub- 
set at the lower end of the scale. Locali- 
ties 5 and 7 are from the same statistical 
population, because they are both in- 
cluded in the second subset. 

Statistical significance must be in- 
terpreted with caution. One cannot 
infer gene pool similarities from knowl- 
edge of a single character; for example, 
it would be improper to consider liz- 
ards from central Mexico and Costa 
Rica as belonging to the same biological 
population, even if they are shown to 
belong to the same statistical popula- 
tion. It is well known that phenotypic 
expression of the genotype can be modi- 
fied by environmental pressures and 
similarities, or differences in a particu- 
lar character among samples from di- 
vergent localities simply may be an 
expression of this pressure on a rela- 
tively plastic genome. 

Sexual Dimorphism 

Presence or absence of sexual dimor- 
phism for individual characters was de- 
termined by analysis of variance. This 
was done for each locality sample of 
Ameiva festiva and undulata, for all 
males against all females of A. ameiva, 
chaitzami, and leptophrys, and for At- 
lantic slope males versus Atlantic slope 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



13 



females and Pacific slope males versus 
Pacific slope females of quadrUineata. 
If any sample as so defined was sevually 
dimorphic for a character, the entire 
If any sample as so defined was sexually 
dimorphic for that character for pur- 
poses of statistical analysis. Presence or 
absence of sexual dimorphism for most 
of the characters utilized in this study is 
indicated in Table 1. 

Sources of Specimens Examined 

A complete list of specimens exam- 
ined, along with locality data for each, 
is included in Appendix B. The follow- 
ing abbreviations have been used in 
referring to individual specimens and 
indicate the museum or private collec- 
tion in which the specimens are housed. 

ANSP Academy of Natural Sciences of 

Philadelphia 

AMNH American Museum of Natural 

History 

ARP A. R. Philips Collection 

(University of Arizona) 



BMNH 


British Museum (Natural History) 


FMNH 


Field Museum of Natural History 


JRM 


John R. Meyer (Private 




Collection) 


LACM 


Los Angeles County Museum 


MCZ 


Museum of Comparative Zoology 


MNHN 


Museum National d'Histoire 




Naturelle, Paris 


RDS 


Richard D. Sage (Private 




Collection) 


TCWC 


Texas Cooperative Wildlife 




Collection 


USNM 


United States National Museum 


UCR 


Universidad de Costa Rica 


UCLA 


University of California at Los 




Angeles (Specimens now 




housed in the Los Angeles 




County Museum) 


UIMNH 


University of Illinois Museum of 




Natural History 


KU 


University of Kansas Museum of 




Natural History 


UMMZ 


University of Michigan Museum 




of Zoology 


USC(CRE) 


University of Southern California 




(Costa Rican Expedition) 


UU 


University of Utah 


ZMB 


Zoologisches Museum, Berlin 



SYSTEMATIC ACCOUNT OF THE SPECIES 

I have recognized six species of KEY TO THE MIDDLE AMERICAN 

Ameiva in Middle America. All of these SPECIES OF AMEIVA 

are easily recognizable in the field by , „ i i . t i r 

the experienced student, but are easily ^- ^^" ^? *^^^^^^ longitudmal rows of 

confused when dealing with preserved ^^^^''''^ /^'"^f^; "^^^optychial scales 

specimens. As is often the case, the subequal; adults spotted _. .. amewa 

species cannot be distinguished readily Eight longitudinal rows of ventral 

on the basis of single characters, and scutes; transverse row of enlarged 

identification of preserved specimens mesoptychial scales; principally 

can be difficult even if a suite of char- striped 2 

acters is used. The following artificial 2. Posterior gular scales smaller than 

key should be used in conjunction with anterior gular scales 3 
the tables in the analyses of geographic 

variation and with the descriptions of Anterior and posterior gular scales 

color and pattern variation. No attempt subequal m size 4 

has been made to place the key into a 3. Postnasal scales not in contact with 

phylogenetic framework. Fortunately, prefrontals; lateral parietal and fron- 

the situation is not so bad as that which toparietal scales separated by one or 

caused Stuart (1955), in discussing his more rows of small, irregular scales; 

key to Guatemalan Anolis, to comment midgular scales greatly enlarged, ir- 

that ". . . the worker who knows what regular in arrangement leptophrys 

species he has before him should ex- Postnasal scales in contact with pre- 

perience few difficulties in its use." frontals; parietal and frontoparietal 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



82° 



78° 




50 100 



KILOMETERS 



82° 



78° 



Fig. 2. Map showing locality records for Ameiva ameiva in Panama. To avoid crowding of sym- 
bols, some localities have been omitted. Samples used in statistical analyses were ( 1 ) Vicinity of 
Panama City, (2) Vicinity of Colon, (3) Vicinity of Niievo Gorgona, (4) Isla San Jose, (5) Prov. 
Herrera, (6) Prov. Los Santos, and (7) Vicinity of Cerro Campana. 



.scales in contact; midgular scales 
slightly to moderately enlarged, ir- 
regular or in a single longitudinal 
row quadrilineata 

Midgular scales much enlarged, ir- 
regular in arrangement; narrow, 
light-colored vertebral stripe present 
except in large adults festiva 

Midgular scales much enlarged, in 
longitudinal arrangement or not, 
moderately enlarged and irregular in 
arrangement; no vertebral stripe . 5 

Small (maximum observed SVL 85 
mm for males, 75 mm for females); 
paravertebral stripe narrow (mean 
PV = 37.1); dorsolateral blotches in 
males fused to dorsolateral light 
stripe chaitzami 

Moderately large (maximum ob- 
served SVL 129 mm for males. 111 
for females ) ; paravertebral 



mm 



stripe broad (mean PV = 47.4 for 
males, 46.4 for females); dorsolateral 
blotches in males, if present, do not 

fuse with dorsolateral light stripe 

undulata 



Ameiva ameiva (Linnaeus) 

Lacerta ameiva Linnaeus, Systema naturae, ed. 
12, 1:362, 1766.— America. 

Cnemidophorus praesignis Baird and Girard, 
Proc. Acad. Nat. Sci. Philadelphia, 6:129, 
1852 [Syntypes: USNM 5519 and UMMZ 
3823, Chagres, Panama; C. B. Adams, col- 
lector]. 

Ameiva praesignis — Cope, Proc. Acad. Nat. 
Sci. Philadelphia, 14:67, 1862. 

Ameiva praesigna — Bocourt, Mission Scienti- 
fique au Mexique et dans I'Amerique Cen- 
trale; fitudes sur les reptiles, livr. 3, p. 265, 
pi. 20b, fig. 9-9d, 1874. 

Ameiva surinamensis — Boulenger, Catalogue of 
the Lizards in the British Museum (Natu- 
ral History), 2nd Ed., vol. 2, p. 352, 1885 
(part). Giinther, Biologia Central!- Ameri- 
cana; Reptilia and Batrachia, p. 22, pi. 19, 
1885 (part). 

Ameiva ameiva praesignis — Barbour and 
Noble, Bull. Mus. Comp. Zool., 59(6):468, 
1915. Dunn, Proc. Acad. Nat. Sci. Phila- 
delphia, 92:113, 1940. 

Distribution in Tanaind. — Grasslands 
and savannas of the Pacific slopes near 
Chepo, Panama, to David, Chiriqui, and 
in open areas across the Canal Zone to 
Colon and Portobelo, Colon, on the 
Atlantic side (Fig. 2). 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



io 



Diagnosis. — Ameiva ameiva can be 
distinguished from all other Middle 
American Ameiva by the following com- 
bination of characters: Size large, max- 
imum observed SVL 197 mm for males 
and 157 mm for females; mesoptychial 
scales small, irregular; scales around the 
tail more than 30; ventrals at midbody 
usually more than eight; supraoculars 
usually eight; color pattern in adult 
males emphasizes spots, rather than 
longitudinal stripes or lateral bars and 
blotches. 

Description. — Maximum observed 
SVL 197 mm for males, 157 mm for 
females; 11-16 (12.9) supralabials; 8-12 
(10.1) infralabials; 6-10 (8.0) supraocu- 
lars; COF values 2-6 (2.1), but note 
geographic variation discussed below; 
COP values 8-16 (10.6); nostril in pre- 
nasal-postnasal suture; prefrontals in 
contact with postnasals; loreals 2; GAB 
128-204 (146.0), 204 an exceptional 
value — the second highest value re- 
corded was 169; GOR 229-324 (287.8) 
for males, 225-394 (295.5) for females; 
GAB/GOR 0.45-0.68 (0.51) for males, 
0.41-0.67 (0.49) for females; SAT 34-45 
(39.5); total preanal scales 7-16 (10.8); 
terminal preanal scales 2-3 (2.4) for 
males, 2-3 (2.2) for females; one row of 
enlarged prebrachials with irregularly 
arranged accessory rows on either side; 
postbrachials slightly to moderately en- 
larged, irregularly arranged; preante- 
brachials enlarged, one (occasionally 
two) rows; prefemoral scales enlarged, 
in four or more rows; infratibial scales 
in three rows; femoral pores 25-41 
(32.4); subdigital lamellae 26-36 (30.9); 
longitudinal rows of ventral scutes 28- 
33 (31.2); transverse rows of ventral 
scutes 6-12 (11.0); dorsal caudal scales 
weakly keeled. For details of scutella- 
tion see figure 3. 

Color and Pattern in Alcohol. — Juve- 
nile males: Top and upper half of side 
of head gray-green to brown with black 
markings immediately anterior to eye; 
lower one-half of head from eye to post- 
nasal white; supralabials black above, 




Fig. 3. Head and preanal scutellation of 
Ameiva ameiva from Panama (KU 108258): 
( A ) Lateral view of head, ( B ) Dorsal view of 
head, (C) Ventral view of head. X L3. 
(D) Preanal region. X 2. 



white below to lip; infralabials black 
below, white above to lip; chin and 
gular region white or cream with small, 
scattered flecks and often a transverse 
row of black flecks between larger me- 
dian gulars and smaller posterior gulars; 
middorsal area between occiput and tail 
black or dark brown with brown or 
gray-green middorsal stripe, anterior 
one-fifth of which as broad as paraver- 
tebral field, gradually narrowing to a 
width of about 10 granules near base of 
tail and with irregular margins through- 
out; cream or gray-white paravertebral 
stripes from (and including) supercili- 
ary series well onto tail; dorsolateral 
dark stripe black with or without a 
brown median stripe and with many 
small, white or blue- white spots; lateral 
light stripe cream, originating at dorso- 
posterior margin of ear and extending 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



to point of insertion of hind leg and 
onto anterior surface of thigh, beginning 
again on posterior surface of thigh and 
extending well onto tail; area between 
lateral light stripe and ventral scutes 
black with small, white spots (some in- 
distinct); venter white or cream medial- 
ly with lateral scutes and (occasionally) 
chest marked with black Hecks; dorsal 
surfaces of fore and hind limbs black 
with blue, blue-white or pale brown 
spots and blotches, this color and pat- 
tern grading to uniform cream or white 
ventrally with or without black Hecks; 
pre- and postfemoral light stripes cream. 
Tail brown with black or blue-gray 
flecking above, flecks sometimes ar- 
ranged to give the appearance of 
whorls; white dorsolateral stripe; sides 
black with white flecking; ventrolateral 
light stripe white, blending into white 
or cream ventral coloring; subcaudal 
area sometimes with black blotches; 
short, black postanal stripes extending 
from each corner of the vent. 

Through ontogeny there is a gradual 
loss of stripes and an increased empha- 
sis on spotting, so that in adult males 
the dorsal pattern is one of transverse 
rows of white or blue-white spots on a 
black background with indistinct gray- 
green or pale brown areas between the 
rows of spots (Fig. 4). Anteriorly, the 
pattern breaks down altogether, leaving 
only fine, light brown reticulations on a 
black background. This pattern carries 
over onto the dorsal surfaces of the 
forelimbs and the granules of the side 
of the head. The hind limbs become 
black with blue flecks, again sometimes 
arranged to give the appearance of 
whorls. Ventrally, the body, tail and 
hind limbs are blue; the chin, gular 
region and forelimbs white or cream 
with black flecks. In adult males the 
chin, gular region, and the forepart of 
the chest may be pale orange. Males 
from eastern Panama retain a bright 
orange or reddish-tan vertebral stripe as 
adults. 

Females: Juvenile females resemble 



juvenile males, but there is strong sex- 
ual dimorphism in adult color patterns 
( Fig. 4 ) . Adult females retain longi- 
tudinal stripes, and spotting does not 
become as pronounced as in adult 
males. Adult females have brownish- 
white to white dorsolateral and lateral 
stripes bounded by black. Middorsally, 
there may be a remnant of the vertebral 
stripe (this is always true of specimens 
from eastern Panama) with the remain- 
ing area mottled gray-green or brown 
on black with scattered white spots. A 
short cream stripe originating at the 
posteroventral corner of the ear and 
ending at the shoulder may be present. 
The dorsolateral dark stripe is black 
with white spots and may contain a dif- 
fuse, brown secondary stripe. Between 
the lateral light stripe and the enlarged 
ventral scutes there is a broad, light 
brown stripe. Some females have no 
spotting at all, the areas between the 
paravertebral and lateral light stripes 
appearing uniformly green or pale 
brown. All combinations of plain and 
spotted patterns in middorsal, dorsolat- 
eral and ventrolateral fields may be 
found. 

Color and Pattern in Life. — The fol- 
lowing description of color in life of 
Atneiva ameiva is quoted (with modi- 
fications to eliminate abbreviations) 
from the field notes of C. W. Myers and 



Fig. 4. Ontogenetic Change and Variation in 
Pattern of Ameiva ameiva from Panama. 
Males: (A) KU 107521, SVL 68 mm, from 
Guanico, Los Santos Prov.; (B) KU 107542, 
107 mm, from Pese, Herrera Prov.; (C) KU 
107532, 160 mm, from Guanico, Los Santos 
Prov. Females: (D) KU 76106, 45 mm, from 
S Slope Cerro Campana, Panama Prov.; (E) 
KU 95546, 92 mm, from the Rio Salado, 
Chiriqui Prov.; (F) KU 107524, 132 mm, from 
Guanico, Los Santos Prov.; (G) KU 108259, 
150 mm, from Juan Mina, Chagres River, 
Canal Zone. Small males resemble small fe- 
males as exemplified by KU 76106 (D above). 
KU 108259 (G above) is included to illustrate 
the presence of a well defined vertebral stripe 
in adult lizards from eastern Panama ( see 
te.xt for discussion). 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



17 




18 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



pertains to specimens collected at 
Guanico Arriba, Los Santos, Panama: 
(KU 107521-3; juveniles) ". . . Mid- 
dorsum brown with darker brown marks 
and bounded by pale yellow dorsolat- 
eral stripe. Sides rich brown or blackish 
brown, with pale tan dots, bordered 
below by a creamy stripe." ( KU 107524- 
7; adult females) ". . . Dorsolateral and 
lateral stripes remain conspicuous, and 
the dorsum takes on a greenish tinge or 
not; a variable amount of tan to yellow 
dorsal spots. Throat light orange, and 
undersides of arms orange tinged to 
light dull red; rest of venter white in 
one and pale blue in the others; rear of 
the thighs white or pinkish white. Iris 
light brown." (KU 107528-31; adult 
males) ". . . Body stripes absent or in- 
conspicuous and body with many con- 
spicuous light spots which tend to be 
greenish yellow above and pale yellow 
on sides. Venter as in females but there 
seems to be a tendency for the colors, 
especially the orange, to be brighter 
(but the belly of one is bluish white). 
Iris light brown." 

Geographic Variation. — Sample lo- 
calities are indicated in figure 2. Of the 
characters analyzed by STP, the follow- 
ing show no among-sample statistical 
significance (ranges of sample means in 
parentheses ) : Total infralabials ( 9.8- 
10.2), total supraoculars (8.0-8.1), COF 
values for females (2.0), GOR (males, 
282.5-295.8; females, 288.5-318.2), and 
GAB/GOR (males, 0.50-0.51; females, 
0.47-0.50). As previously stated, A. 
ameiva from eastern Panama differ from 
individuals from western Panama in 
their retention of a distinct vertebral 
stripe as adults. Specimens collected 
west of a line between Penonome, 
Code, and the mouth of the Rio Estan- 
cia, Code, lack the stripe as adults. 

On the basis of STP results, the 
sample from Isla San Jose in the Bay of 
Panama (Sample 4) can be distin- 
guished from other samples in three 
characters (within sample ranges fol- 
lowed by sample means in parenthe- 



ses): Total supralabials 14-15 (14.4), 
other samples 11-16 (12.3-12.8); COF 
values for males 2-6 (4.5), other sam- 
ples 2 (2.0); COP values 8-12 (9.2), 
other samples 8-15 (10.0-12.0). Addi- 
tionally, the Isla San Jose sample has 
the largest sample means for the fol- 
lowing characters, although the sample 
does not differ statistically from main- 
land samples for these characters: total 
supraoculars, GAB, GOR for females, 
GAB/GOR for males, terminal preanals 
for females, and transverse rows of ven- 
trals. The sample values were lowest 
among the localities compared for GAB/ 
GOR for females, femoral pores, ven- 
trals at midbody and COP values. With 
respect to the total number of supra- 
labials, the significant difference be- 
tween sample 4 and other samples is 
probably a function of the few large 
samples available for analysis. That in- 
clusion of other samples in the compari- 
son might alter the picture is suggested 
by values obtained for this character 
from small, mainland samples and from 
small samples from other islands in the 
Bay of Panama. Values intermediate 
between that obtained for Isla San Jose 
and mainland samples used in STP 
analysis were obtained from Isla Taboga 
(mean = 13.0, n = 5); Rio Mamoni, 
Panama (13.6, 10); vicinity of Nata, 
Code (13.6, 5); Rio Chorcha east of 
Chiriqui, Chiriqui (13.8, 5); and Isla 
San Miguel (14.0, 6). A sample of five 
specimens from Tres Puntas, Herrera, 
has a mean of 14.6, higher than that of 
the Isla San Jose sample for total num- 
ber of supralabials. 

Samples 5 and 6 from the Azuero 
Peninsula can be distinguished statisti- 
cally as a unit from other samples for 
COP values, 12 (12.0) and 10-13 (11.9), 
respectively; other samples 8-15 (9.2- 
10.5). These samples possess the great- 
est (although not statistically signifi- 
cant) mean values for total number of 
femoral pores and number of subdigital 
lamellae. 

On the mainland of Panama, distinct 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



19 



geographic trends are apparent only for 
the number of femoral pores, number of 
termhial preanals of males and COP 
values. All of these characters show in- 
creasing values from east to west. 
Values obtained for small samples not 
used in STP analysis indicate the same 
trends. Other characters demonstrate 
no geographic trends although certain 
localities differ from others statistically. 
Ecological Notes. — Ameiva ameiva 
is characteristic of open areas in Pana- 
ma. It is common in the savannas of the 
lower Pacific slopes and in cleared areas 
across the Canal Zone. Cochran ( 1946 ) 
reported it in the same habitat on Isla 
San Jose. It has adapted well to the 
presence of man and is common in resi- 
dential parts of Panama City. Heatwole 
(1966) reported A. ameiva to be absent 
in the Darien Gap and noted that the 
Panamanian population of the species is 
disjunct from South American popula- 
tions. He postulated that the species 
was once continuously distributed over 
the isthmus but that post-Columbian 
reforestation isolated the Panamanian 
population. Heatwole further demon- 
strated that A. ameiva is presently ex- 
panding its range eastward in Panama 
by moving into areas recently cleared 
by man. 

Remarks. — It is difficult to explain 
the presence of two geographically dis- 
tinct color types (presence or absence 
of a reddish vertebral stripe in adults) 
of Ameiva ameiva in Panama. Present 
environmental conditions are similar 
east and west of the Penonome-Rio 
Estancia line. Bennett ( 1968 ) provided 
a clue to solution of the problem with 
his suggestion that the eruption of Vol- 
can El Valle, which may have occurred 
during the Pleistocene (Terry, 1956), 
was an event of biogeographic im- 
portance. The ejecta from El Valle may 
have covered many square miles and 
could have divided the Panamanian 
population of Ameiva ameiva for suf- 
ficient time for divergence of color pat- 
tern to occur. Detailed ecological in- 



vestigations of A. ameiva might yield 
infomiation on this point, as well as on 
the origins of east-west clines for certain 
other characters, but no such studies 
have been undertaken. Even statistical 
studies aimed at correlating geographic 
variation in characters with environ- 
mental factors cannot be carried out 
until samples of adequate size for analy- 
sis can be accumulated from through- 
out the range of the species in Panama. 
Taxonomic recognition of segments of 
the Panamanian population of Ameiva 
ameiva would be premature until analy- 
sis of character variation throughout the 
range of the species in South America 
can be completed. 

Two specimens of Ameiva suritia- 
mensis ('=A. ameiva) were listed by 
Boulenger (1885:353) as collected in 
Costa Rica by "Mr. Geale." There are 
no records at the British Museum that 
these lizards (BMNH 80.6.21.1-2) were 
collected by Ceale himself, although the 
museum bought a number of specimens 
from him between 1866 and 1880 (Mr. 
A. F. Stimson, pers. comm. ) . The speci- 
mens are indeed A. ameiva and can be 
referred to the subspecies praesignis 
which ranges through northwestern 
Venezuela, northeastern Colombia and 
Panama, but it is doubtful that they 
were collected in Costa Rica. Both 
specimens possess relatively well-de- 
fined middorsal stripes, suggesting that 
they may have come from eastern 
Panama, if they were collected in Cen- 
tral America at all. There is presently 
no suitable habitat for A. ameiva in 
Costa Rica with the possible exception 
of the dry Guanacaste region where 
the species is not now known. 

Barbour and Noble ( 1915 ) examined 
specimens of Ameiva ameiva ostensibly 
from Acapulco, Mexico. Re-examina- 
tion of the specimens (MCZ 2728 and 
2730) confirms the identification, but 
the locality information is obviously 
erroneous. 

Yunker and Radovsky ( 1966 ) and 
Brennan and Yunker ( 1966 ) reported 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 




10' 



8' 



50 100 

I -I 



KILOMETERS 



Fig. 5. Map showing locality records of Ameiva Jeptophnjs. To avoid crowding of symbols, some 
localities have been omitted. Samples used in statistical analyses were ( 1 ) Osa Peninsula, Punta- 
renas Prov., Costa Rica, (2) Palmar-Golfito Region, Puntarenas Prov., Costa Rica, (3) Barro Colo- 
rado Island, Canal Zone, Panama, and (4) Eastern Darien Prov., Panama. 



the acarinids Dracomjsus belgicae 
Yunker and Radovsky, Eutrombicula 
alfreddugesi (Oudemans) and E. goeldi 
(Oudemans) parasitic on Ameiva hi- 
frontata from Panama. Examination of 
the series of Uzards from which these 
parasites were taken proves them to be 
A. ameiva. 

Ameiva leptophrys Cope 

Amiva leptophrijs Cope, Proc. Amer. Phil. Soc, 
31:341-342, 1893 [Holotype: Apparently 
lost; given by Cope as "No. 318." Type 
Locality: "Buenos Ayres" (= Buenos Aires, 
Puntarenas Prov., Costa Rica)]. 

Ameiva leptophrys — Dunn, Proc. Acad. Nat. 
Sci. Philadelphia, 92:114, 1940. Stuart, 
Proc. Biol. Soc. Washington, 55:146, 1942. 
Taylor, Univ. Kansas Sci. Bull., 38(1): 
257-260, 1956. 

Ameiva nithveni Barbour and Noble, Bull. 
Mus. Comp. Zool., 59(6) :471-473, 1915 
[Holotype: MCZ 10927. Type Locality: 
near Panama City. Collector: W. W. 
Brown, Jr.]. Burt and Burt, Trans. Acad. 
Sci. St. Louis, 28(1 ):53, 1933. 

Ameiva festiva — Wettstein, Sitzungsli. Akad. 
Wiss. Wien, math.-natiuw. Kb, Abt. I, 143. 
Bd., 1.U.2 Heft. p. 30, 1934 (part). 

Distribution. — Forested areas of 
Pacific slopes from Parrita, Puntarenas 



Prov., and San Isidro del General, San 
Jose Prov., Costa Rica to Jaque, Darien 
Prov., and the Rio Mono, Darien Prov., 
within a few kilometers of the Colom- 
bian border in Panama; across the 
Canal Zone and east along the Atlantic 
slopes at least as far as the Rio Sasardi, 
San Bias Prov., Panama (Fig. 5). The 
species probably occurs in extreme 
northwestern Colombia. 

Diagnosis. — Ameiva leptophrys can 
be distinguished from its Middle Amer- 
ican congeners by the following com- 
bination of characters: Separation of 
parietal and frontoparietal scales from 
the interparietal by one or more rows of 
small, irregular scales; abrupt reduction 
in size of small scales immediately pos- 
terior to the enlarged, median gular 
scales; large number of femoral pores 
(mean number for males, 49.1; females, 
44.7); a transverse row of abruptly en- 
larged mesoptychial scales; eight longi- 
tudinal rows of ventral scutes; color 
pattern emphasizing longitudinal stripes 
rather than spotting, but lacking a light 
vertebral stripe. 

Description. — Maximum observed 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



21 




Fig. 6. Head and preanal scutellation of 
Arneiva Icptophrijs: (A) Lateral view of head, 
KU 96827, (BJ Dorsal view of head, KU 
96827, (C) Ventral view of head, KU 96827. 
X 1.2. (D) Preanal region, KU 96815. X 1. 

SVL 133.0 for males, 129.0 for females; 
supralabials 11-15 (12.5) for males, 12- 
15 (12.7) for females; infralabials 10-12 
(10.1); supraoculars 6-8 (6.2); COF 
values 2-6 (3.5); COP values 5-10 
(8.1); SO-SC values 2-10 (5.9); loreals 
2-6 (3.7); nostril in prenasal-postnasal 
suture; prefrontal usually not in contact 
with postnasal (96.3%, N = 136); GAB 
127-182 (153.8); PV 28-63 (41.0); PV/ 
GAB 0.19-0.35 (0.26); GOR 182-285 
(235.6); GAB /GOR 0.54-0.74 (0.66); 
SAT 18-25 (21.0); total preanal Scales 
4-10 (6.6) for males, 6-10 (8.1) for fe- 
males; terminal preanal Scales 1-3 (2.1) 
for males, 1-3 (1.9) for females; one 
row of enlarged prebrachials; postbra- 
chials not enlarged to moderately en- 
larged, irregularly arranged or in one or 
two rows; preantebrachials enlarged, in 



two rows proximally, one row distally; 
3 (usually) or 4 rows of enlarged pre- 
femoral scales; two rows of enlarged 
infratibials; femoral pores 42-61 (49.1) 
for males, 34-53 (44.7) for females; sub- 
digital lamellae 24-35 (28.1); longi- 
tudinal rows of ventral scutes 26-31 
(28.8); transverse rows of ventral scutes 
8-10 (8.1). For details of scutellation 
see figure 6. 

Individual variation in dorsal head 
scutellation is extreme (Fig. 7). There 
is no predictable pattern to scutellation 
posterior to the frontal, and even this 
scale may be divided. However, usually 
two frontoparietal and three parietal 
scales can be recognized. The degree 
of variation in posterior head scales is 
unequalled by other species of Middle 
American Ameiva, although it is ap- 
proached in the closely related A. fes- 
tiva. 




Fig. 7. Variation of dorsal head scutellation of 
Ameiva leptophrys. (A) KU 96820, X 2, 
(B) KU 96826, X 2.2, (C) KU 96814, X 2.3, 
(D) KU 95577, x 3.8, (E) KU 25641, x 4.1. 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



Color and Pattern in Alcohol. — Juve- 
nile males: Top of head light brown; 
rostral and anterior part of frontonasal 
pearl-white with a slight greenish tinge; 
side of head dark brown above grading 
to gray-brown below; area between 
orbit and ear brown with darker brown 
postorbital bar dorsally; white crescent 
beneath and slightly behind eye; chin, 
infralabials, gular region and throat im- 
maculate. Middorsal stripe broad, 
slightly irridescent, gray-brown, bor- 
dered laterally by white paravertebral 
stripes originating on superciliary scales 
and extending well onto tail; paraverte- 
bral stripes (and edges of middorsal 
stripe) undulating on posterior three- 
quarters of body; velvet-black dorso- 
lateral stripe from above shoulder to 
insertion of hind legs, bordered below 
by broken white line originating on 
shoulder and extending to insertion of 
hind limbs and onto anterior surface of 
femur; area below this lateral light 
stripe brown, grading to gray-brown 
above ventral scutes. Venter immacu- 
late. Dorsal surfaces of fore and hind 
limbs dark brown with lighter brown 
blotches, spots or mottling; no post- 
femoral light stripe; ventral surfaces of 
limbs immaculate to gray-white. Tail 
brown with black Hecks above, brown 
laterally (below dorsolateral light 
stripe) blending to white or gray- white 
ventrally; short postanal white stripe 
bounded by brown below for about six 
scales, then joining white ventral pat- 
tern. 

Through ontogeny ( Fig. 8 ) the mar- 
gins of the middorsal stripe become 
more undulating and the paravertebral 
stripes are lost except where the mid- 
dorsal stripe remains broadest. Gray- 
brown or dark brown bars or large spots 
appear laterally overlaying the dorso- 
lateral and ventrolateral dark fields. In 
some large adult males the lateral light 
stripes are lost entirely, and the venter 
becomes blue-white or coppery from 
chin to vent and onto the ventral sur- 
face of the tail. 



Females: Juvenile females are simi- 
lar to juvenile males. Through ontogeny 
( Fig. 8 ) the middorsal stripe often be- 
comes interrupted by narrow, dark 
brown cross-bars. The adult pattern of 
females is similar to that of adult males, 
faint to relatively distinct bars appear- 
ing in the dorsolateral and ventrolateral 
dark fields. However, some females lack 
lateral bars, and the lateral light stripes 
are retained throughout life. 

Color and Pattern in Life. — The dor- 
sal coloration in life of Ameiva lepto- 
phry.s does not differ markedly from 
that seen in preserved specimens. The 
middorsal stripe is metallic brown and 
may have a greenish tint. Paravertebral 
and lateral light stripes are yellowish to 
yellowish-gray, and lateral bars or 
blotches, if present, are yellowish-gray 
to rust colored. Ventral coloration is 
variable. The entire ventral surface may 
be white, cream or copper colored in 
juvenile males and females. Adults of 
both sexes may retain these colors, but 
some adult males become pale blue 
over the entire ventral surface, or they 
may have greenish-yellow, orange or 
brick red on chin, gular region and 
throat with the ventral surfaces of body, 
limbs and tail blue. Some adult males 
exhibit a variation of the latter color 
pattern in which the hind limbs pos- 
terior to the femoral pores and the 
ventral surface of the tail is copper- 
colored. Variability in ventral colora- 



FiG. 8. Ontogenetic Change and Variation in 
Pattern of Anieiva Jeptophnjs. Males: (A) 
KU 107555, SVL 85 mm from N slope Cerro 
Cambutal, Los Santos Prov., Panama; (B) KU 
76142, 115 mm from Barro Colorado Island, 
Canal Zone, Panama; (C) KU 95582, 118 mm 
from 6 km W Palmar Norte, Puntarenas Prov., 
Costa Rica (included for comparison with KU 
76142). Females: (D) KU 95578, 44 mm 
from 6 km W Palmar Norte, Puntarenas Prov., 
Costa Rica; (E) KU 76144, 89 mm from Barro 
Colorado Island, Canal Zone, Panama; (F) 
KU 76163, 112 mm from El Valle, Code Prov., 
Panama. Small males are similar to small 
females as exemplified by KU 95578 ( D 
above ) . 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



23 




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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



tion might be correlated with reproduc- 
tive condition in these Hzards, but this 
aspect of their biology has not been 
investigated. It \\'ould be instructive to 
retain specimens in captivity in order to 
ascertain not only when color changes 
occur, but whether or not greenish-yel- 
low and orange pigmentation represent 
stages leading to brick red coloration. 

Geographic Variation. — Any analysis 
of geographic variation of characters of 
Ameiva leptophnjs must be considered 
preliminary, pending the acquisition of 
additional large samples of specimens 
from throughout the range of the 
species. Only four such samples have 
been available for this study (Fig. 5). 
There is a noticeable lack of specimens 
from western Panama, and the follow- 
ing results should be viewed with both 
the paucity of large series of specimens 
and the lack of good geographic repre- 
sentation in mind. 

The following characters exhibit no 
significant geographic differences among 
sample means (ranges of sample means 
in parentheses): Supralabials (males, 
12.1-12.9; females, 12.4-13.2), infra- 
labials (10.0-10.3), SO-SC values (5.2- 
6.3) and terminal preanal scales of 
males (1.9-2.3). 

Samples 1 and 2 (Costa Rica) are 
significantly different from samples 3 
and 4 (Panama) with respect to the 
following characters (locality number 
followed by sample mean in parenthe- 
ses): Number of subdigital lamellae 
(Sample 1, 29.2; 2, 29.4; 3, 26.5; 4, 27.2), 
paravertebral granules (1, 45.9; 2, 48.9; 

3, 38.1; 4, 37.4) and PV/GAB (1, 0.28; 
2, 0.30; 3, 0.25; 4, 0.25). Samples 1, 2 
and 3 differ significantly from sample 4, 
but not from each other, in mean COF 
values (1, 7.4; 2, 7.3; 3, 8.0; 4, 8.7). One 
character, total number of femoral 
pores, shows a significant difference be- 
tween samples 1 and 2 ( 1, 43.6; 2, 48.4 ) . 
In addition to COF values, mentioned 
above, sample 3 differs significantly 
from sample 4 in GAB/GOR (3, 0.68; 

4, 0.65), but sample 4 is not significantly 



different from samples 1 and 2 (both 
0.66). There are a number of charac- 
ters for which mean values of either 
sample 1 or 2 differ significantly from 
that of sample 3, but not from sample 
4, and a few (COP, GAB, Longitudinal 
Rows of Ventrals) for which mean 
values of sample 1 and/ or sample 2 are 
not significantly different from sample 
3, but do differ from the mean value for 
sample 4. 

East-west clines are apparent for a 
number of characters. In terms of mean 
values, there is a decrease from east to 
west ( infralabials, longitudinal rows of 
ventrals) or an increase from east to 
west (COF values, terminal preanals of 
females, total preanals of females). If 
samples 1 and 2 are combined (it is 
legitimate to do so according to STP 
results), additional characters show the 
same clinal tendencies. An increase in 
mean value from east to west is noted 
for number of supralabials of males, 
number of supraoculars, GAB, PV and 
PV/GAB; decreasing mean values from 
east to west are noted for COP values. 
A small sample (n =r 10) of leptophnjs 
from near the Rio Sasardi in San Bias 
Prov., Panama, was collected by C. W. 
Myers and arrived too late to be in- 
cluded in the STP analysis. Mean 
values calculated for each character for 
the Rio Sasardi sample are in accord 
with the east-west trends discussed 
above for (Rio Sasardi means in paren- 
these): COF values (2.8), total pre- 
anal scales of females (8.4), number of 
supralabials of males (12.2) and COP 
values (9.1). The Rio Sasardi sample 
runs counter to east-west trends estab- 
lished on the basis of mean values of 
samples 1-4 for number of terminal pre- 
anals of females (2.0), number of in- 
fralabials (10.0), longitudinal rows of 
ventrals (28.7), number of supraoculars 
(6.3), GAB (160.1) and PV/GAB 
(0.24). These results once again call 
attention to the necessity of obtaining 
additional samples from intermediate 
localities before attempting a really 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



25 



definitive analysis of geographic varia- 
tion of leptophrys. 

With the possible exception of color 
pattern, none of the characters not ana- 
lyzed by STP appears to vary geo- 
graphically. Male leptophrys from cen- 
tral and eastern Panama tend to have 
more emphasis on spotting (rather than 
bars) in the lateral dark fields then do 
specimens from western Panama and 
Costa Rica. With the acquisition of 
more specimens from the hiatus in west 
and central Panama this difference may 
also prove to be clinal. In addition, 
specimens from the Osa Peninsula of 
Costa Rica ( sample 1 ) appear darker in 
overall coloration than those collected 
at other localities, including those from 
the adjacent non-peninsular mainland of 
Costa Rica. Since my sample from the 
Osa Peninsula was collected over a 
short period of time and by the same 
collectors, it may be that the darkness 
of the specimens can be attributed to 
preservation in warm formalin or to an 
unusually long period of time in fomia- 
lin before transfer to ethyl alcohol. 

Ecological Notes. — Amewa lepto- 
phrys is a forest species; at various lo- 
calities throughout its range it is found 
sympatrically with all other species of 
Middle American Ameiva, except chait- 
zami. Hillman (1969) studied niche 
specificity of leptophrys, festiva and 
quadrilineata near Rincon on the Osa 
Peninsula of Costa Rica. He found that 
adult leptophrys forage deeper into the 
forest than its sympatric congeners; the 
latter search for food in more open 
areas. Roth leptophrys and festiva seem 
to utilize the same areas for basking, 
but there is little overlap with the more 
heliophilous quadrilineata. Hillman 
found hatchling leptophrys occupying 
more open areas than the adults but 
virtually identical to areas utilized by 
adult festiva. Presumably the size dif- 
ference between juvenile leptophrys and 
adult festiva acts to lessen direct com- 
petition. In the vicinity of Palmar 
Norte, Puntarenas Prov., Costa Rica, 



festiva is not abundant, and I found 
leptophrys and quadrilineata more 
closely associated ecologically than 
either is to festiva. The two species 
were seen basking within a few meters 
of one another, but whereas quadriline- 
ata utilized clearings, leptophrys basked 
in the sun-flecked areas beneath the 
dense, low vegetation. Ameiva lepto- 
phrys would allow approach to within 
a meter before bolting when so hidden. 
Tree-fall clearings in a swamp forest 
near Palmar Norte were occupied by 
juvenile quadrilineata and leptophrys, 
but I never found the young of both 
species utilizing the same clearing. 
Dunn (1940a) found leptophrys and 
festiva sympatric on Rarro Colorado 
Island but reported that festiva oc- 
cupied the more shaded parts of the 
habitat, whereas leptophrys utilized 
more open areas, the opposite of the 
situation in Costa Rica. Charles W. 
Myers (personal communication) found 
leptophrys and ameiva sympatric at 
Guanico Arriba, Los Santos Prov., Pana- 
ma. There leptophrys occupies dry, 
partly disturbed forest and ameiva is 
common in open pastures. Recently col- 
lected material from near Parrita, Pun- 
tarenas Prov., Costa Rica, confirms the 
sympatry of leptophrys and undulata 
hesitantly reported by Dunn (in Stuart, 
1942). 

Remarks. — Rarbour and Loveridge 
(1929b: 213) listed two paratypes of 
Ameiva ruthveni (^=A. leptophri/s): 
MCZ 10925-10926. Rarbour and Noble 
(1915:471-472) acknowledged examina- 
tion of only two specimens in their de- 
scription of A. ruthveni: MCZ 9931 (re- 
catalogued MCZ 10927; E. E. Williams, 
in. litt.), the holotype, and MCZ 9932 
(recatalogued MCZ 10926). The third 
hzard, MCZ 10925 (originally MCZ 
9928), is not a paratvpe. One paratype 
(MCZ 10926) was not located. 

Brennan and Yunker (1966:235-236, 
258) reported Ameiva undulata from 
Panama as host for the acarinids Eu- 
tromhicula alfreddugesi (Oudemans) 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



and E. goeldi (Oudemans). Examina- 
tion of the specimens of Ameiva prove 
them to be leptophrys. There are no 
vahd records of A. undiihta from 
Panama. 

Ameiva festiva (Lichtenstein and 
Von Martens) 

Cnemidophorus festivus Lichtenstein and Von 
Martens, Nomenclator Reptilium et Am- 
philiiorum Musei Zoologici Berolinensis, p. 
13, 1856 [Lectotype, herein designated: 
ZMB 881a; Paralectotype, formerly syn- 
type: ZMB 881b. Type Loca]it>-: "Vera- 
gua" = Veragiias, Panama]. 

Ai7iewa eutropia Cope, Proc. Acad. Nat. Sci. 
Philadelphia, 14:62, 1862 [Syntypes: 
ANSP 9071, USNM 4320(3). Type lo- 
cality: "Region of the Triiando, New Gre- 
nada" (=Colombia). Collector: Arthur 
Schott]. 

Aiyjeiva edwardsi Bocourt, Ann. Sci. Nat. 17 
(17): 1-2, 1873 [Holotype: MNHN 5480. 
Type Locality: Izabel and Santa Maria de 
Panzos near Rio Polochic, Guatemala. Col- 
lector : unknown] . 

Ameiva festivus — Bocourt, Mission Scientifique 
au Mexique et dans TAmerique Centrale; 
fitudes sur les reptiles, livr. 3, pp. 260-263, 
pi. 20A, Fig. 10; pi. 20D, Fig. 3; pi. 20, 
Fig. 2; 1874. 

Ameiva festiva — Giinther, Biologia Centrali- 
Americana; Reptilia and Batrachia, p. 24, 
1885. Boulenger, Catalogue of the Lizards 
in the British Museum (Natural History), 
vol. 2, p. 347, 1885. Barbour and Noble, 
Bull. Mus. Comp. Zool., 59(6):473-4, 
1915. Burt and Burt, Bull. American Mus. 
Nat. Hist., 61(5) :309, 1931. Burt and 
Burt, Trans. Acad. Sci. St. Louis, 28(1): 
53, 1933. Wettstein, Sitzungsb. Akad. 
Wiss. Wien, math.-naturw. Kl., Abt. I, 143. 
Bd., l.u. 2 Heft., p. 30, 1934. 

Ameiva festiva festiva — Dunn, Proc. Acad. 
Nat. Sci. Philadelphia, 92:114, 1940. Tay- 
lor, Univ. Kansas Sci. Bull., 38(1):260- 
264, 1956. 
Ameiva festiva edwardsi — Stuart, Occ. Pap. 
Mus. Zool., Univ. Michigan, No. 471, p. 
21, Figs. 6-7, 1943. Smith and Laufe, 
Univ. Kansas Sci. Bull. 31(2):64-66, 1945. 
Smith and Taylor, U. S. Nafl. Mus. Bull, 
199:174, 1950. Stuart, Misc. Publ. Mus. 
Zool., Univ. Michigan, No. 122, p. 77, 
1963. 
Ameiva festiva occidentalis Taylor, Univ. Kan- 
sas Sci. Bull., 38(l):260-264, 1956 [Holo- 
type: FMNH 120236. Type Locality: 8.0 
km E San Isidro del General, San Jose 
Prov., Costa Rica. Collector: Richard C. 
Taylor] . 



Distribution. — Ameiva festiva is 
found in the forests of the Caribbean 
lowlands from Tabasco, Mexico to 
Depto. Boyaca, Colombia and on the 
Pacific lowlands from the vicinity of 
San Isidro del General, San Jose Prov., 
Costa Rica, at least to the Condoto- 
Andagoya area in the Intendencia de 
Choco, Colombia. The species also is 
known from the vicinity of Tilaran, 
Guanacaste Prov., Costa Rica, just west 
of the continental divide (Fig. 9). 

Diagnosis. — Ameiva festiva can be 
distinguished from other Middle Amer- 
ican Ameiva by the following combina- 
tion of characters: Central gular scales 
much enlarged, irregular in arrange- 
ment; gradual reduction in size of scales 
radiating outward from enlarged, cen- 
tral gular scales; many femoral pores 
(mean for males, 40.8; females, 40.0); 
a transverse row of abruptly enlarged 
mesoptychial scales; eight longitudinal 
rows of ventral scutes; a distinct, light 
colored vertebral stripe present in all 
but large adults. No other species of 
Ameiva from Middle America possesses 
such a well-defined vertebral stripe, al- 
though several South American Ameiva 
do have such a stripe (edracantha, or- 
cesi, bridgesi, and septemlineata). 

Description. — Maximum obsei"ved 
SVL 114.0 mm. for males, 129 mm. for 
females; supralabials 12-18 (14.2) for 
males, 12-17 (13.9) for females; infra- 
labials 8-15 (10.8); sublabials 4-9 (4.7) 
for males, 4-7 (4.5) for females; supra- 
oculars 4-10 (6.5); COF values 2-6 
(2.6); COP values 2-14 (9.4); loreals 
2; nostril in prenasal-postnasal suture; 
prefrontals in contact with postnasal 
(41.1%) or no (54.5%), 4.5 percent in 
contact on one side but not the other 
(n = 448); GAB 145-221 (182.5) for 
males, 143-213 (179.8) for females; VS 
5-29 (12.8) for males, 4-27 (11.2) for 
females; VS/GAB 0.02-0.14 (0.06); 
COR 209-324 (261.1) for males, 215- 
312 (259.5) for females; GAB/GOR 
0.57-0.85 (0.70); SAT 16-22 (18.3); 
total preanal scales 4-10 (6.6) for males, 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



27 




Fig. 9. Map showing locality records of Aineiva festiva. To avoid crowding of symbols, some 
localities haxe been omitted. Samples used in statistical analyses were ( 1 ) Piedras Negras, Depto. 
El Peten, Guatemala, (2) Vicinity of Chinaja, Depto. Alta Verapaz and Depto. El Peten, Guate- 
mala, (3) Finca Chama, Depto. Alta Verapaz, Guatemala, (4) North Coast Localities, Honduras, 
(5) Bonanza, Depto. Zelaya, Nicaragua, (6) Cukra and Kanawa, Depto. Zelaya, Nicaragua, (7) 
Rio Mico and vicinity, Depto. Zelaya, Nicaragua, (8) Tuli Creek, Depto. Rio San Juan, Nicaragua, 
(9) Puerto Viejo, Heredia Prov., Costa Rica, (10) Turrialba, Cartago Prov., Costa Rica, (11) 
Tortuguero, Limon Prov., Costa Rica, (12) Los Diamantes and vicinity, Limon Prov., Costa Rica, 
(13) Almirante, Bocas del Toro Prov., Panama, (14) Isla Escudo de Veraguas, Panama, (15) 
Barro Colorado Island, Canal Zone, Panama, (16) Rio Mono, Darien Prov., Panama, (17) Golfito 
Region, Puntarenas Prov., Costa Rica, (18) Vicinity of Rincon, Osa Peninsula, Puntarenas Prov., 
Costa Rica, (19) Rio Ure and vicinity, Depto. Cordoba, Colombia, (20) Condoto-Andagoya-Tado 

Area, Intendencia de Choco, Colombia. 



4-9 (6.4) for females; terminal preanal 
scales 1-3 (2.0) for males, 1-3 (2.2) for 
females; one row of enlarged prebra- 
chials; postbrachials enlarged, in a 
single short row; preantebrachials en- 
larged, two rows proximally, one dis- 
tally; three rows of enlarged prefemoral 
scales; two rows of enlarged tibial 
scales; femoral pores 33-55 (40.8) for 
males, 32-50 (40.0) for females; sub- 



digital lamellae 23-34 (28.4); longitu- 
dinal rows of ventral scutes 25-33 
(27.8); transverse rows of ventral scutes 
6-8 (8.0). 

Individual variation in posterior 
head scutellation is great, second only 
to that of Ameiva leptophrijs, but two 
frontoparietal and three parietal scales 
are usually recognizable (Fig. 10). 

Color and Pattern in Alcohol. — 



28 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 




Fig. 10. Head and preanal scutellation of 
Ameiva festiva (ZMB 881a; Lectotype): (A) 
Lateral view of head, ( B ) Dorsal view of 
head, (C) Ventral view of head. X 1.5. (D) 
Preanal region. X 1-6. 

There is extensive geographic variation 
in color and pattern over the range of 
Ameiva festiva. As a basis for discuss- 
ing this variation, the following com- 
plete description of color and pattern 
of specimens from Puerto Viejo, Here- 
dia Prov., Costa Rica, is presented. 

Juvenile males: Blue-gray vertebral 
stripe 5-13 granules wide at midbody 
extending from rostral scale well onto 
tail, the margins slightly scalloped on 
the posterior three-fourths of body; top 
of head lateral to vertebral stripe dark 
gray; side of head dark gray above be- 
coming lighter below, anterior to eye; 
ear encircled by interrupted white line; 
chin, infralabials, gular region and 
throat light gray or white; dorsal sur- 
face of body very dark brown lateral to 
vertebral stripe; dorsolateral light stripe 
white, narrow and interrupted, originat- 
ing on canthus and extending onto tail; 
dorsolateral dark stripe velvet black, 



beginning immediately posterior to eye 
and extending onto tail, bordered below 
by white, narrow, interrupted ventro- 
lateral light stripe originating beneath 
eye, passing over ear to the groin, 
thence onto anterior surface of femur at 
juncture of dorsal granular scales and 
enlarged ventral scales; vertical bars ex- 
tending ventrally from ventrolateral 
light stripe immediately anterior and 
posterior to forelimbs; ventrolateral 
field dark brown with irregular blue- 
gray spots or narrow, regularly spaced 
blue-gray bars, or both. Ventral sur- 
faces of body, limbs and tail gray-white. 
Dorsal surface of forelimbs and femur 
dark brown to black; shank dark brown 
with blue-gray spots above. 

Through ontogeny (Fig. 11) the 
vertebral stripe becomes sinuous and 
less well defined. It may be lost entirely 
on the head, posterior part of the body 
and tail. A light-colored rostral scale is 
retained, as usually is the vertebral 
stripe on the anterior third of the body. 
The head becomes uniformly brown 
above and the pale markings above and 
below the eye, on the side of the neck 
and around the ear are lost. The velvet- 
black dorsolateral dark stripe is re- 
tained. In larger individuals, the mid- 
dorsal area immediately lateral to the 
position occupied by the vertebral 
stripe becomes regularly marked with 
dark and light brown blotches, the 
darker occupying positions where the 
sinuous or scalloped margin of the ver- 
tebral stripe curved inwards toward the 
midline; this gives the back a ladder- 
like pattern. The dorsal surface of the 
tail becomes imiformly brown or brown 
with darker brown blotches which may 
be paired or nearly so. The dorsolateral 
and ventrolateral light stripes are re- 
tained posteriorly only. Most males be- 
come blue-gray overall ventrally, the 
lateral ventral scutes and enlarged fem- 
oral scales marked with black. The in- 
fralabials may also be flecked with 
black. 

Females: The color and pattern of 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



29 





Fig. 11. Ontogenetic Pattern Change of Ameiva f estiva from the Atlantic 

Slopes of Costa Rica. Males: (A) AMNH 89210, SVL 43 mm; (B) AMNH 

89191, 80 mm; (C) AMNH 89203, 106 mm. Females: (D) AMNH 89212, 

47 mm; (E) AMNH 89204, 69 mm; (F) AMNH 89193, 90 mm. 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



juvenile females is similar to that of 
juvenile males. Females differ from 
males in that the juvenile color and pat- 
tern is retained with little fading in all 
but very large adults and adult females 
do not develop blue colors ventrally. 

Color and Pattern in Life. — The fol- 
lowing notes refer to specimens col- 
lected by me on 1 August 1965 at Puerto 
Viejo, Heredia Pro v., Costa Rica. Adult 
male (KU 95558), SVL 99 mm: Neck 
and chest reddish-orange; chin and 
gular region pale blue; belly pale blue 
medially and pale blue broken by black 
blotches laterally; middorsal area brown 
with indications of a faint yellow verte- 
bral stripe extending from occiput to 
base of tail; dorsolateral dark field 
black, bordered above and below by 
narrow, interrupted yellow stripes; ven- 
trolateral field olive with narrow yellow 
bars. Adult female (KU 95559), SVL 
79 mm: Venter pale bronze; dorsal pat- 
tern similar to KU 95558 except verte- 
bral stripe more pronounced and ex- 
tending from rostral well onto tail; yel- 
low on head and body becoming pale 
blue on tail. 

Juvenile Ameiva festiva from near 
Bluefields, Depto. Zelaya, Nicaragua 
(KU 101214-101215) possess vertebral 
stripes which are yellow anteriorly be- 
coming blue-green on the tail. The dor- 
solateral and ventrolateral light stripes 
are yellow to yellowish-orange. An 
adult female (KU 101213) from the 
same locality has the same color and 
pattern as KU 95559 from Puerto Viejo. 

As in other species of Ameiva I have 
examined, gular coloration varies in 
adult males. Collections are not such 
that this can be correlated with season. 
I suspect the color to be indicative of 
reproductive state, at least in males, but 
lack evidence to substantiate this hy- 
pothesis. Field notes taken by W. E. 
Duellman, C. W. Myers, and myself, as 
well as color photographs provided by 
D. H. Janzen disclose that the gular 
region and chest of adult male festiva 
may be uniformly blue (if so, the entire 



venter is blue), copper, dull brick-red, 
yellow-orange, golden-yellow or yellow- 
green (all commonly associated with a 
blue venter, although individuals hav- 
ing a copper gular region often have the 
entire venter that color). There seems 
to be no geographic pattern associated 
with these colors or none that I can de- 
tect from the field notes available to 
me. The ventral surfaces of females 
may be uniformly copper or cream 
(often with a pinkish tint), or may be 
one of these colors with the gular region 
and chest gray. 

Geographic Variation. — Only the 
number of transverse rows of ventral 
scutes and the number of terminal pre- 
anals of males show no significant dif- 
ferences among sample means. A num- 
ber of characters exhibit variation 
among samples, but show no marked 
geographic trends. These characters 
are: GOR, GAB, number of femoral 
pores, number of longitudinal rows of 
ventrals, total number of preanal scales 
and SAT (although SAT values from 
samples in the Atlantic lowlands may 
tend to be higher than values from 
Pacific samples). 

A striking aspect of geographic vari- 
ation among samples of festiva is the 
apparent "center" of variation for cer- 
tain characters in the Caribbean low- 
lands of Costa Rica and Nicaragua. 
Values for VS, VS/GAB, and number of 
sublabials of males gradually increase 
in samples to the north and east of these 
lowlands. A decrease in number of 
supralabials and in COF values is ap- 
parent in the same directions. 

A clinal increase in mean values 
from north (Guatemala) to southeast 
( Panama and Colombia ) is expressed 
for GAB/GOR, but samples 17 (Golfito 
Region of Costa Rica) and 18 (Osa 
Peninsula of Costa Rica) have high 
means, nearer to values noted for Sam- 
ples 16 (Rio Mono, Darien Prov., Pana- 
ma) and 20 ( Condoto-Andagoya-Tado 
Area, Intendencia de Choco, Colombia) 
to the south and east. The mean value 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



31 



for sample 19 ( Rio Ure and vicinity, 
Depto. Cordoba, Colombia) is lower 
than would be predicted by the trend. 
A clinal decrease from north to south- 
east is expressed in number of infra- 
labials ( Samples 1 and 19 have lower 
and higher mean values, respectively, 
than anticipated by the trend), number 
of sublabials of females (although high 
values are noted for samples 14 and 20 ) , 
COP values (locality 20 has a very low 
mean and represents a separate subset) 
and the number of subdigital lamellae. 
As is evident, these clinal tendencies are 
never perfectly expressed: one or more 
samples disrupt the trend. Samples 19 
and 20 (in South America) counter in 
some respects geographic trends estab- 
lished in the more northerly parts of the 
range of festiva. These two localities 
are separated by high mountains and 
differ from one another in a statistically 
significant way for a number of charac- 
ters. Samples 15 (Barro Colorado Is- 
land) and 16 (Rio Mono, Darien Prov., 
Panama) both show greater similarities 
to Sample 20 than to Sample 19. Sam- 
ples 17 and 18, in southeastern Costa 
Rica, show similarities to Sample 19 for 
some characters, to sample 20 for others. 
It would be instructive to have addi- 
tional samples from the Pacific slopes 
of western Panama in order to see 
whether there is further evidence for 
the "Panamanian crossover" suggested 
by Dunn (1940b). 

The more northerly samples of fes- 
tiva (Samples 1-4) differ from others in 
a number of characters, none of which 
set these samples apart statistically from 
all others. The samples from Guatemala 
(Samples 1-3) and Honduras (Sample 
4), taken as a group, differ from adja- 
cent samples in a statistically significant 
way for number of supralabials of males 
and females, VS of males (data from 
Sample 4 unavailable) and total pre- 
anal scales of females. 

As alluded to earlier, there is marked 
geographic variation in the color and 
pattern over the range of festiva. There 



are four distinct pattern types with 
variation to a lesser degree within two 
of these. The four patterns are dis- 
cussed separately below relative to the 
description of specimens from Puerto 
Viejo, Costa Rica, given above. 

Pattern Type A: The color and pat- 
tern described for festiva from Puerto 
Viejo is characteristic in major aspects 
of specimens collected in the Caribbean 
lowlands of Costa Rica, western Pana- 
ma, and northward into Mexico. Liz- 
ards comprising Samples 1-7 and pos- 
sibly 8 (Nicaragua northward) tend to 
be browner in overall coloration than 
those from Costa Rica and Panama 
(Samples 9-13), although this is a sub- 
jective observation and not based upon 
quantitative evidence. The first impres- 
sion one gets upon seeing living festiva 
from Costa Rica and Panama is one of a 
black lizard with a prominent vertebral 
stripe. The lizards from Honduras and 
Guatemala have a broader vertebral 
stripe than others with Pattern Type A; 
this is especially striking in specimens 
from Guatemala. 

Pattern Type B: A second widely 
distributed pattern type, largely allo- 
patric to Pattern Type A, is peculiar to 
lizards from the Pacific slopes of south- 
eastern Costa Rica and western Pana- 
ma, the San Bias Coast of Panama and 
Colombia east of the Cordillera Central 
(Samples 15-19). Specimens from this 
area are characterized by a relatively 
broader vertebral stripe than that of 
other pattern types and by having the 
ventrolateral light stripe either lacking, 
present or present only posteriorly. The 
vertebral stripe broadens posteriorly and 
is broadest in specimens from eastern 
Panama and Colombia. Lizards from 
the San Bias Coast of Panama and from 
east of the Cordillera Central in Colom- 
bia possess vertebral stripes that may 
become as broad as the middorsal area 
at the base of the tail (Fig. 12B). 
Ameiva festiva from the Osa Peninsula 
of Costa Rica (Sample 18) are darker 
in overall coloration than those from the 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 





Fio. 12. Patterns of Ameiva f estiva from Colombia. (A) FMNH 165748, male, SVL 44 mm, from 
the Upper Rio Ure, Depto. Cordoba. (B) FMNH 165227, male, 101 mm, from the Upper Rio 
Ure, Depto. Cordoba. (C) FMNH 43828, female, 81 mm, from Pizzaro, Intendencia de Choco (?). 



adjacent mainland. Thi.s is not the re- 
sult of preservation, for color photo- 
graphs taken by Daniel H. Janzen be- 
fore the lizards were preserved show 
them to be darkly pigmented. Two 
specimens of f estiva ( KU 80210 and KU 
110745) from Almirante, Bocas del Toro 
Prov., Panama, exhibit a pattern similar 
to that described here, although others 
from the same locality show Pattern 
Type A. Ameiva festiva from Barro 
Colorado Island and southeastern Pana- 
ma are somewhat intermediate between 
Pattern Types A and B, but because the 
population more closely resembles the 
latter, I have included them here. 

Pattern Type C: Ameiva festiva 
from the Choco of Colombia (including 
Sample 20) have a pattern similar to 
Type A, but the vertebral stripe is ex- 
tremely straight and well defined even 
in large specimens ( Fig. 12C ) . The 
dorsolateral and ventrolateral light 
stripes are also well defined, may be 
entire or dotted (rarely dashed), and 
are not lost in large individuals. Pre- 
served specimens are, with the excep- 



tions of the light stripes and vertebral 
stripe, very dark. 

Pattern Type D: Adult Ameiva fes- 
tiva from Isla Escudo de Veraguas, off 
the northwest coast of Panama, are 
melanistic. That this is so is supported 
by color photographs of freshly col- 
lected specimens provided by Charles 
W. Myers. Very little pattern is ap- 
parent on specimens I have examined 
and, although examination of juvenile 
specimens (unavailable to me) might 
prove them to belong to Pattern Type 
A or B, I prefer to assign them to a 
separate pattern type for the present. 
Baskin and Williams (1966) summa- 
rized information on insular populations 
of melanistic Ameiva in the Caribbean, 
and Mayr (1963) discussed similar 
populations of Lacerta in the Mediter- 
ranean. Such melanistic island popula- 
tions are usually found on small, diy, 
barren islands ( Mayr, 1963 ) ; Heatwole 
and Torres ( 1966 ) attributed the phe- 
nomenon in Ameiva to local diftcrentia- 
tion, those species of Ameiva exhibiting 
melanism not being closely related. 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



33 



Mayr ( 1963 ) suggested that the dark 
pigmentation might function in heat 
regulation. The sample from Isla Es- 
cudo de Veraguas docs not fit the pat- 
tern in temis of habitat as outlined 
above. According to C. W. Myers (per- 
sonal communication), one of the few 
biologists to have visited the island, Isla 
Escudo de Veraguas ". . . is covered 
with humid forest, including swamp 
forest, except for a well developed line 
of strand vegetation at some points." 
Specimens of festiva from the Bocas del 
Toro Archipelago to the west of Isla 
Escudo de Veraguas are not notably 
mclanistic and conform to Pattern Type 
A, although, according to Myers, habi- 
tats on all of these islands are similar. 
That the melanism of festiva on Isla 
Escudo de Veraguas is the result of 
local differentiation can hardly be 
denied, but the adaptive significance of 
such pigmentation remains a mystery. 

Ecological Notes. — In the northern 
part of its range, Ameiva festiva in- 
habits heavy forest (Stuart, 1943; Smith 
and Laufe, 1946; Echternacht, 1968) 
when sympatric with other macroteiids. 
If other large teiids are not present, the 
niche of the species is expanded to in- 
clude more open habitats (Echternacht, 
1968). In Mexico, Guatemala, and 
Honduras, festiva is partly sympatric 
with undulata and with various species 
of Cnemidophorus. In Costa Rica and 
Panama, festiva and either leptophnjs 
or quadrilineata, or all three species 
may occur together. In the vicinity of 
Almirante and on the islands of the 
Bocas del Toro Archipelago, Bocas del 
Toro Pro v., Panama, festiva and quad- 
rilineata are sympatric, as they are on 
Isla Escudo de Veraguas. Ameiva fes- 
tiva typically occupies the more shaded 
parts of the habitat, whereas quadriline- 
ata occurs in the open (C. W. Myers, 
personal communication). In southeast- 
ern Costa Rica (vicinity of San Isidro 
del General, San Jose Prov.; vicinity of 
Palmar Sur and on the Oso Peninsula, 
Puntarenas Prov.) festiva is sympatric 



with both quadrilineata and leptophnjs. 
Ameiva festiva occupies a habitat in- 
termediate to those of the other two 
species: leptophnjs occupies more 
shaded areas and quadrilineata, the 
open areas. This has been noted by 
Hillman (1969) and myself, but Dunn 
( 1940a ) reported the habitats of lepto- 
phnjs and festiva reversed on Barro 
Colorado Island in Panama; there fes- 
tiva occupies the less shaded areas. 
Hillman ( 1969 ) found hatchling A. lep- 
tophnjs sympatric with adult festiva on 
the Osa Peninsula and concluded that 
the difference in size between the two 
minimizes competition. The reproduc- 
tive biology of festiva has been investi- 
gated by R. E. Smith (1968a, b) in the 
only such study of Ameiva that has been 
undertaken. Smith's results suggest that 
festiva is reproductively active through- 
out the year, and that clutch sizes aver- 
age between 2.22 and 2.29 eggs. 

Nothing has been reported on the 
ecology of A. festiva in South America. 

Remarks. — Ameiva festiva was de- 
scribed by Lichtenstein and von Mar- 
tens ( 1856 ) on the basis of two speci- 
mens, which are catalogued under one 
number (ZMB 881) in the Zoological 
Museum of Berlin. The description is 
based on a specimen "Long, a rostro ad 
caud. bas. 3 1/2", caud. ultra 6"" (88.9 
mm). The syntypes presently have 
snout- vent lengths of 86 and 58 mm; the 
tail of the larger is broken. I am herein 
designating the larger syntype as the 
lectotype; shrinkage resulting from over 
one hundred years in preservative would 
account for the discrepancy in snout- 
vent length between that given in the 
description and my measurement. The 
smaller syntype becomes a paralecto- 
tyi^e. 

Dunn (1943) described Ameiva fes- 
tiva nicefori from Saisima, Depto. Cun- 
dinamarca, Colombia, but I have pre- 
sented evidence elsewhere ( Echter- 
nacht, 1970) that the population de- 
serves specific recognition. Dunn (1940a) 
and Villa (1968) allied the population 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 




Fig. 13. Map showing locality records of Ameiva quadrilineata. To avoid crowding of symbols, 
some localities have been omitted. Samples used in statistical analyses were ( 1 ) Southeastern Nica- 
ragua, (2) Tortuguero, Limon Prov., Costa Rica, (3) Limon and vicinity, Limon Prov., Costa Rica, 
(4) Bocas del Toro Prov., Panama, (5) Isla Escudo de Veraguas, Panama, (6) San Isidro del 
General, San Jose Prov., Costa Rica, (7) Palmar Sur and vicinity, Puntarenas Prov., Costa Rica, 
(8) Vicinity of Rincon, Osa Peninsula, Puntarenas Prov., Costa Rica, (9) Progreso, Puerto Ar- 
muelles and vicinity, Chiriqui Prov., Panama, (10) Boquete, Chiriqui Prov., Panama. 



of Ameiva on the Islas del Maiz, Nica- 
ragua with undulata rather than festiva 
as originally described (Barbour and 
Loveridge, 1929a). My investigations 
support this conclusion, and the popu- 
lation is discussed with undulata in the 
present work. 

Ameiva quadrilineata (Hallowell) 

Cnemidophorus quadrilineatus Hallowell, Proc. 
Acad. Nat. Sci. Philadelphia, 1860, p. 483, 
1860 [Syntypes: USNM 6053A, 6053B. 
Type Locality: "Nicaragua," restricted to 
Greytown, Depto. Rio San Juan, by Tay- 
lor, Univ. Kansas Sci. Bull., 38(1):271, 
1956]. 



Ameiva quadrilineata — Cope, Proc. Acad. Nat. 
Sci. Philadelphia, 1862, p. 61, 1862. Bo- 
court, Mission Scientifique au Mexique et 
dans TAmerique Centrale, fitudes sur les 
Reptiles, livr. 4, p. 259, pi. 20A, Fig. 9, 
1874. Giinther, Biologia Centrali-Ameri- 
cana, Reptilia and Batrachia, p. 24, 1885. 
Dunn, Proc. Acad. Nat. Sci. Philadelphia, 
92:113, 1940. Taylor, Univ. Kansas Sci. 
Bull., 38(1):271, 1956. 

Ameiva <^ahhiana Cope, J. Acad. Nat. Sci. 
Philadelphia, ser. 2, 8:33, pi. 28, Fig. 3, 
1875 [Syntypes: USNM 32614-16. Type 
Locality: Old Harbor (= Puerto Viejo), 
Limon Prov., Costa Rica. Collector: W. M. 
Gabb]. Giinther, Biologia Centrali-Ameri- 
cana, Reptilia and Batrachia, p. 23, 1885. 

Ameiva undulata — Boulenger, Catalogue of 
the Lizards in the British Museum (Na- 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



35 




Fig. 14. Head scutellation of Ameiva quad- 
lilineata: (A) Lateral view of head, (B) Dor- 
sal view of head, (C) Ventral view of head; 
KU 95593 from 1.5 km E Palmar Norte, 
Puntarenas Prov., Costa Rica. X 2. (D) Dor- 
sal view of head; KU 96849 from Cayo 
Zapatillo Grande, Bocas del Toro Prov., 
Panama, X 2.7. 



tiiral History), 2nd Ed. 
1885 (part). 



vol. 2, p. 348, 



Distribution. — Atlantic coastal low- 
lands from extreme southeastern Nica- 
ragua to northwestern Panama; Pacific 
slopes of Costa Rica from Quepos, Pun- 
tarenas Prov. (H. S. Fitch, personal 
communication) to the Azuero Penin- 
sula, Panama; the Bocas del Toro Archi- 
pelago, Isla Escudo de Veraguas and 
Isla de Cebaco of Panama and Isla del 
Cafia of Costa Rica. Specimens have 
been collected at elevations of 1100- 
1650 m above Paso Ancho and on Cerro 
Hornito, Chiriqui Prov., Panama (Fig. 
13). _ 

Diagnosis. — A small lizard which 
can be distinguished from other Middle 
American Ameiva by the following com- 
bination of characters: Central gular 
scales moderately enlarged, irregular, 




hJ^^S 



Fig. 15. Preanal region of Ameiva quadrili- 
ncata: (A) KU 95636 from 6 km W Palmar 
Norte, Puntarenas Prov., Costa Rica, X 2. 
(B) KU 95593 from 1.5 km E Palmar Norte, 
Puntarenas Prov., Costa Rica, X 2. 



scales of posterior one-half of gular re- 
gion abruptly smaller than scales of 
anterior one-half (Figs. 14-15); moder- 
ate number of femoral pores (mean 
number for males, 34.2; females, 27.8); 
a transverse row of abruptly enlarged 
mesoptychial scales; eight longitudinal 
rows of ventral scutes; narrow middor- 
sal field (mean PV: 26.1) which does 
not become darker laterally; continu- 
ous, well defined dorsolateral stripe and 
continuous to interrupted ventrolateral 
stripe, both usually retained throughout 
life. The middorsal field of Ameiva un- 
diilata is broad (mean PV: males, 46.8; 
females, 45.6) and the ventrolateral 
light stripe is usually interrupted. 
Ameiva chaitzami also has a relatively 
broad middorsal field (mean PV: 38.1) 
and interrupted ventrolateral light 
stripes and, in addition, lacks a uni- 
formly colored middorsal field. 

Description. — Maximum observed 
SVL 88 mm for males, 82 mm for fe- 
males; supralabials 11-16 (13.7); infra- 
labials 10-13 (10.1); supraoculars 6-9 
(6.7); COF values 2-6 (3.0); COP 
values 4-12 (9.1); loreals 2-4; nostril in 
prenasal-postnasal suture; prefrontals in 
contact with postnasals; GAB 116-177 
(140.5); PV 16-36 (26.1); PV/GAB 
0.09-0.25 (0.19); GOR 185-258 (218.6); 
GAB/GOR 0.41-0.78 (0.64) for males, 
0.56-0.75 (0.64) for females; SAT 17-22 
(19.8) for males, 17-21 (19.5) for fe- 
males; total preanal scales 4-10 (7.1) 
for males, 4-9 (6.8) for females; termi- 
nal preanal scales 1-3 (1.4) for males, 



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ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



37 



1-3 (1.3) for females; prebrachials en- 
larged, in a single row; postbrachials 
moderately enlarged, in one row or ir- 
regular; preantebrachials in three rows 
proximally, one or two rows distally; 
three rows of enlarged prefemoral 
scales; infratibials enlarged, in two 
rows; femoral pores 22-42 (34.2) for 
males, 18-39 (27.8) for females; sub- 
digital lamellae 25-34 (29.5) for males, 
25-34 (28.8) for females; longitudinal 
rows of ventral scutes 28-33 (29.8); 
transverse rows of ventral scutes 4-10 
(8.0). 

Color and Pattern in Alcohol. — Juve- 
nile males: Top of head light brown to 
gray-brown; side of head gray with 
black wash above grading to gray on 
lower half of postnasals, loreals and 
supralabials; middorsal field dark brown 
to dark gray-brown, narrow, bounded 
laterally by narrow, white dorsolateral 
light stripes, which originate above the 
eyes and extend to the base of the tail 
where they coalesce and form a verte- 
bral stripe extending well onto the tail; 
dorsolateral dark fields black; ventro- 
lateral light stripe narrow, white, orig- 
inating beneath eye and extending to 
the groin, thence onto anterior surface 
of femur where it runs between the dor- 
sal granular scales and the ventral 
plates; short, narrow, interrupted white 



<1 

Fig. 16. Ontogenetic change and variation in 
pattern of male Ameiva quadrilineata. A-C; 
Pacific population. D-G; Atlantic population. 
Females do not show marked ontogenetic 
changes in color or pattern and resemble 
juvenile males as to these characters. (A) KU 
95623, SVL 31 mm, from the Rio Zapote, 8 
km E Palmar Norte, Puntarenas Prov., Costa 
Rica; (B) KU 95583, 68 mm, from 8 km SE 
San Isidro del General, San Jose Prov., Costa 
Rica; (C) KU 93984, 84 mm, from 2-5 km 
ESE Piedras Blancas, Puntarenas Prov., Costa 
Rica; (D) KU 96837, 40 mm, from Isla Basti- 
mentos, Bocas del Toro Prov., Panama; (E) 
KU 108282, 62 mm, from E end Isla Escudo 
de Veraguas, Panama; (F) KU 96845, 79 
mm, from Cayo Zapatillo, Bocas del Toro 
Prov., Panama; (G) Ventral view of KU 96845 
to show dark pigmentation. 



stripe from beneath ear to shoulder; 
ventrolateral dark field black with 
brown mottling below; dorsal surfaces 
of forelimbs mottled brown and black, 
an interrupted white line extending 
along the back of the arm from shoulder 
to wrist; dorsal surface of hind limbs 
mottled black and brown with white 
spots on shank and narrow, white post- 
femoral stripe extending onto tail for 
ca. 10-12 scale whorls; ventral surfaces 
immaculate except for faint gray mark- 
ings in the gular region and a faint, 
transverse gray line across the gular re- 
gion at the juncture of the large anterior 
and small posterior scales. 

Through ontogeny (Fig. 16) black 
mottling appears in the middorsal area, 
but never enough to disrupt the visual 
impression of a unicolored field. Gray 
or blue-gray reticulations appear in the 
ventrolateral dark field, this pattern ex- 
tending onto the lateral-most enlarged 
ventral scutes and the enlarged ventral 
femoral scales. The dorsolateral dark 
field remains black above, but brown 
vertical bars or reticulations appears in 
the lower half. The white spots on the 
hind limbs and the short stripe between 
ear and shoulder disappear, the side of 
the neck becoming uniformly gray- 
brown or black with gray reticulations. 
Ventral surfaces may be pale blue over- 
all or the chin, gular region and throat 
may be salmon colored, the remainder 
of the venter pale blue. There may be 
black flecks in the midventral region 
(see discussion of geographic varia- 
tion ) . 

Females: Juvenile females resemble 
juvenile males. Adult females tend to 
retain the juvenile pattern except that 
black flecking appears in the middorsal 
field and small, brown, longitudinally 
oriented flecks and blotches may appear 
in the dorsolateral dark field (Fig. 16). 

There is some geographic variation 
in color and pattern and the foregoing 
description applies to specimens from 
the Pacific lowlands of Costa Rica. 

Color and Pattern in Life. — ^The fol- 



38 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



lowing description is extracted from the 
field notes of C. W. Myers and refers to 
a series of Ameiva quadriUneata (KU 
96853-96865) collected 13 km N Puerto 
Armuelles, Chiriqui Prov., Panama, on 
22 April 1965. Ameiva quadriUneata 
from this locality do not differ in color 
and pattern from individuals found in 
the Pacific lowlands of Costa Rica, the 
latter having been used for the descrip- 
tion of color and pattern in alcohol. 
The following account (modified from 
Myers' field notes) sei-ves to illustrate 
the variability of ventral pigmentation. 
Dorsolateral and lateral stripes vivid 
yellow in small individuals and some- 
what duller in larger ones; dorsolateral 
stripes nearly disappearing in largest 
males. Middorsal region light yellow- 
brown or gray-brown, with or without 
irregular black spotting, and turning to 
a whitish-gray or pale brown dorsal 
stripe on basal one-third of tail. Venter 
of tail base copper, but all but basal 
part of tail is uniformly brown or gray 
above and below (in a juvenile male 
this tail is light blue-gray). Dorsolat- 
eral area of body (between yellow 
stripes) bright rust, variably suffused 
with black (in some nearly uniformly 
black with rust spots). Flanks light 
brown or light gray-brown, heavily 
spotted with black. Ventral color vari- 
able, as follows: KU 96853-5—3 juve- 
niles ( at least two are males ) : Ventral 
surfaces pale copper. KU 96856-9 — 4 
females: Ventral surfaces pale copper, 
perhaps with a yellow tinge on throat. 
KU 96860-1—2 males: Underside of 
head pale blue; rest of venter light blue, 
except tail base and thighs behind fem- 
oral pores, which are light copper. KU 
96862-3—2 males: Underside of head 
light coppery orange; rest of venter 
light blue, except for copper thighs be- 
hind femoral pores, and light blue and 
copper-mottled tail base. KU 96864-5 — 
2 males: Underside of head red-orange 
(pale in smaller specimen and deep in 
larger) with an occasional speck or spot 
of blue showing through; rest of venter 



light blue, except rear of thighs, and 
tail base (copper). 

In addition to the ventral colors 
given above, both males and females of 
all sizes may have immaculate or pearly- 
white ventral surfaces. The chin, gular 
region and throat may be various shades 
of yellow. 

Geographic Variation. — The range of 
Ameiva quadriUneata is divided by the 
high mountains of the Cordillera de 
Talamanca of Costa Rica and Panama. 
Although the resulting Atlantic and Pa- 
cific populations do not show statisti- 
cally significant differences for any char- 
acters, it is apparent that divergence 
has taken place. Mean values for num- 
ber of femoral pores of females and 
number of terminal preanals of both 
sexes are higher and lower, respectively, 
on the Atlantic slope than on the Pacif- 
ic. The mean number of terminal pre- 
anal scales of females from Boquete, 
Chiriqui Prov., Panama (Sample 10), is 
statistically different from mean values 
of all other samples and constitutes a 
separate subset. Several other charac- 
ters show similar trends, but with one 
sample running counter to the trend 
established by the others. Characters 
having lower values along the Atlantic 
slope are GAB, GAB/GOR for both 
sexes, and total number of preanal 
scales for both sexes. The sample from 
Boquete, which is intermediate in geo- 
graphic position to Puerto Armuelles 
(Pacific slope) and Almirante (Atlantic 
slope) in Panama, has intermediate 
mean values to those of Puerto Armu- 
elles and Almirante for some characters 
and more closely resembles samples 
from the Atlantic slope than those from 
the Pacific slope for other characters. 
Boquete lies on the Pacific slope of the 
Cordillera de Talamanca but a few 
kilometers from the continental divide 
in western Panama. Sample 10 has in- 
termediate values for number of supra- 
labials, PV, and SAT of males. Mean 
values for number of supraoculars, 
GAB, GAB/GOR of males and females, 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



39 



and total number of preanals of both 
sexes are nearer mean values for Atlan- 
tic samples than for Pacific samples. 
Occasionally, other samples on the 
Pacific slope resemble the aggregate of 
Atlantic slope samples in terms of mean 
values of one character or another, but 
none with such noticeable frequency as 
does Sample 10. 

Samples 8, 9 and 10 show a statis- 
tical relationship to one another for sev- 
eral characters and constitute a separate 
subset for COF and COP values. Sam- 
ples 8 and 9 on a separate subset for 
GAB, having very high means for this 
character. For GAB, Sample 10 has the 
lowest observed mean value and is simi- 
lar to Atlantic slope samples in this. 
Sample 6 has the lowest observed mean 
PV and constitutes a separate subset. 

General clinal tendencies are noted 
for some characters. The total number 
of preanal scales decreases from north 
to south on the Atlantic slopes, but in- 
creases from north to south in samples 
from the Pacific slopes (except Sample 
10, which has a very low mean); COP 
values decrease from north to south on 
the Pacific side; COF values increase 
from north to south on the Pacific slope; 
the number of subdigital lamellae of 
females decreases from north to south 
on the Atlantic slope. Inclusion of more 
samples in the analysis is necessary in 
order to substantiate these trends. 

Mean values for total number of in- 
fralabials and for numbers of rows of 
longitudinal and transverse ventral 
scutes show no significant differences 
among the ten samples. 

Some geographic variation is appar- 
ent in the color and pattern of Ameiva 
quadrilineata. Although specimens 
from Atlantic and Pacific slope popula- 
tions do not differ markedly among 
themselves, there are differences be- 
tween the two populations. Adult males 
from the Atlantic side are never as 
ornate in terms of dorsal and lateral 
color and pattern as those from the 
Pacific slopes (Fig. 16). Males from 



the Atlantic slopes, especially those 
from northwestern Panama (including 
Samples 4 and 5) have all ventral sur- 
faces variably flecked with black (Fig. 
16); the dorsolateral light stripes of 
specimens from this area may be scal- 
loped over the posterior third of the 
body and may become diffuse, blending 
with middorsal coloration, in large in- 
dividuals; the lateral light stripe may 
be interrupted and the dorsal caudal 
stripe is less evident; the lateral ground 
color of some adults is black mottled 
with brown above, gray below. In gen- 
eral, it seems that the chin, gular region 
and throat of adult male quadrilineata 
from the Atlantic slopes becomes yel- 
low when brightly pigmented, but that 
such lizards from the Pacific slopes 
show red or orange coloration. I know 
of only one exception to this: KU 95600 
collected at Quebrada Boruca at the 
Rio Puente Nuevo, 22 km E Palmar 
Norte, Puntarenas Pro v., Costa Rica, on 
15 August, 1965, was yellow on the 
ventral surface of the head. This gener- 
ality needs to be substantiated; I have 
had no color-in-life notes on Atlantic 
slope specimens from west of the Bocas 
del Toro region of Panama. 

Ameiva quadrilineata from Isla Es- 
cudo de Veraguas of Panama and from 
the Osa Peninsula of Costa Rica do not 
appear to be any darker in coloration 
than those from the adjacent mainlands. 

Ecological Remarks. — Ameiva quad- 
rilineata is ecologically the best known 
member of the genus ( Hirth, 1963, 1965; 
R. E. Smith, 1968a, b). Both Hirth and 
Smith worked with populations of quad- 
rilineata on the Atlantic lowlands, Costa 
Rica. Hirth (1963) studied all aspects 
of the ecology of A. quadrilineata at 
Tortuguero, Limon Prov., Costa Rica. 
At Tortuguero these lizards were abun- 
dant on the beaches, which they shared 
with Basiliscus vittatus. Ameiva quad- 
rilineata is also characteristic of the 
beach fauna on the islands of the Bocas 
del Toro Archipelago and Isla Escudo 
de Veraguas of Panama (C. W. Myers, 



40 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



personal communication) and near the 
northern (western) Hmit of its range 
on the Pacific coast of Costa Rica ( H. S. 
Fitch, personal communication). I have 
collected it in open areas (clearings in 
forested areas, along roadsides, etc.) 
near San Isidro del General and Palmar 
Norte in southeastern Costa Rica. Dvmn 
( 1940a ) reported the species to be com- 
mon in open areas along large rivers on 
the Atlantic side of Costa Rica. Hill- 
man (1969:477), who studied quadrili- 
neata on the Osa Peninsula of Costa 
Rica, found them foraging "in low vege- 
tation adjacent to an exposed surface 
such as a road." The species is defi- 
nitely more heliophilic than either fes- 
tiva or leptophrys, with which it is 
sympatric in some areas. It is similar in 
habitat preference to ameiva. The rela- 
tively small size of adult qiiadrilineata 
undoubtedly helps to reduce competi- 
tion between it and its larger, sympatric 
congeners. Competition may be rela- 
tively more intense between juvenile 
qiiadrilineata and juvenile leptophrys 
or festiva. I have no experience with 
festiva on this point, but I found juve- 
niles of the other two species common 
in tree-fall clearings in a swamp forest 
near Palmar Norte in Costa Rica. The 
clearings were always occupied by one 
of the two, but not both species. 

R. E. Smith (1968a, 1968b) studied 
Ameiva qiiadrilineata and festiva at 
Pandora, Limon Pro v., Costa Rica. His 
investigations indicate that both species 
may be reproductively active through- 
out the year. The mean clutch size for 
qiiadrilineata is 2.05-2.07 eggs, depend- 
ing upon method of counting, and ovi- 
position occurs between 17 and 21 days 
following ovulation. Smith also dis- 
cussed the adaptive significance of fat 
bodies and provided experimental evi- 
dence for a relationship between fat 
body development and gonadal cycles. 

Ameiva are usually diurnal, but 
W. E. Duellman (personal communica- 
tion) found an adult male quadrilineata 
active at night on a road in western 



Panama. There had been a heavy rain 
earlier in the day, and possibly the liz- 
ard was flooded from its burrow. 

Henry S. Fitch (personal communi- 
cation) is studying a population of 
quadrilineata near Quepos, Puntarenas 
Pro v., Costa Rica. The nearest locality 
from which specimens of undulata are 
available is near Parrita, about 25 km to 
the northwest, but Fitch has observed 
both species at Quepos. There quad- 
rilineata occurs only on the beach, and 
undulata is inland in plantations or pas- 
tures where shade and ground cover are 
abundant. Quepos is the northern-most 
locality for quadrilineata on the Pacific 
coast, and the southern-most locality for 
undulata. 

Remarks. — Barbour and Noble 
( 1915 ) apparently confused Ameiva 
quadrilineata and A. undulata pulchra. 
Both were described by Hallowell 
( 1860 ) from "Nicaragua," but quad- 
rilineata is restricted to the Atlantic 
coast of Nicaragua and pulchra to the 
Pacific coast; Barbour and Noble had 
before them specimens from "Chinan- 
dega, Nicaragua," which is on the Pacif- 
ic side. Moreover, the color description 
given by Barbour and Noble fits that of 
undulata, not quadrilineata. 

Ameiva undulata (Wiegmann) 

Cnetnidophorus undulatus Wiegmann, Herpe- 
tologica Mexicana, pp. 27-28, 1834 [Syn- 
types: ZMB 867-869, 869 missing. Type 
Locality: "Mexico," restricted to Tehiian- 
tepec, Oaxaca, Mexico by Hobart M. Smith, 
Proc. Biol. Soc. Washington, 53:56, 1940. 
Collector: Unknown]. 

Ameiva undulatus — Gray, Catalogue of the 
lizards in the British Museum ( Natural 
History), p. 20, 1845. 

Ameiva undulata — Cope, Proc. Acad. Nat. Sci. 
Philadelphia, 14:62-63, 1862. Bocourt, 
Mission Scientifique an Mexique et dans 
TAmerique Centrale; fitudes sur les rep- 
tiles, livr. 3, pp. 254-259, pi. 20A, Fig. 7a- 
7e, pi. 20B, Fig. 1. Gunther, Biologia Cen- 
trali-Americana; Reptilia and Batrachia, 
pp. 23-24, 1885. Boulenger, Catalogue of 
the lizards in the British Museum ( Natural 
History), vol. 2, pp. 347-348, 1885. Hart- 
weg and Oliver, Occas. Papers Mus. Zool., 
Univ. Michigan, 359:7-8, 1937. Dunn, 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



41 



» 



Proc. Acad. Nat. Sci. Philadelphia, 92:114- 
115, 1940. 

Ameiva pulchra Hallowell, Proc. Acad. Nat. 
Sci. Philadelphia, p. 483, 1860 [Syntypes: 
ANSP 9133-9134. Type Locality: "Nica- 
ragua." Collector: Unknown]. 

Cnemidophorus amivoides Cope, Proc. Acad. 
Nat. Sci. Philadelphia, pp. 198-199, 1894 
[Holotype: AMNH 16316. Type Locality: 
La Carpintera, San Jose Prov., Costa Rica. 
Collector: A. Alfaro]. 

Ameiva undulaia undulata — Barbour and 
Noble, Bull. Mus. Comp. Zool, 59(6):474- 
475, 1915. Stuart, Proc. Biol. Soc. Wash- 
ington, 55:145, 1942. Smith and Laufe, 
Univ. Kansas Sci. Bull., 31(2):62-64, Fig. 
2b, 1946. Smith and Taylor, U. S. Natl. 
Mus. Bull., 199:174, 1950. 

Ameiva undulata parva Barbour and Noble, 
Bull. Mus. Comp. Zool., 59(6) :476-477, 
1915 [Holotype: MCZ 5831. Type Locali- 
ty: "Guatemala," restricted to Mazatenan- 
go, Depto. Suchitepequez by Smith and 
Laufe, Univ. Kansas Sci. Bull., 31(2) :51, 
1946. Collector: Unknown]. Stuart, Proc. 
Biol. Soc. Washington, 55:145, 1942. 
Smith and Laufe, LTniv. Kansas Sci. Bull., 
31(2):51-54, Fig. la. Pi. la, 1946. Smith 
and Taylor, U. S. Natl. Mus. Bull., 199: 
173, 1950. Taylor, Univ. Kansas Sci. Bull., 
38(1):26S-271, 1956. Stuart, Misc. Publ. 
Mus. Zool., Univ. Michigan, 122:77, 1963. 

Ameiva undulata pulchra — Dunn, Proc. Acad. 
Nat. Sci. Philadelphia, 92:115, 1940. 
Stuart, Proc. Biol. Soc. Washington, 55: 
146, 1942. 

Aineiva undulata haitwegi Smith, Proc. Biol. 
Soc. Washington, 53:55, 1940 [Holotype: 
USNM 108600. Type Locality: Across the 
Rio Usumacinta from Piedras Negras, 
Depto. El Peten, Guatemala. Collector: 
H. M. Smith]. Stuart, Proc. Biol. Soc. 
Washington, 55:145, 1942. Smith and 
Laufe, Univ. Kansas Sci. Bull., 31(2):35- 
36, Pi. lib, 1946. Smith and Taylor, U. S. 
Natl. Mus. Bull., 199:171-172, 1950. 
Stuart, Misc. Publ. Mus. Zool., Univ. Mich- 
igan, 122:77, 1963. 

Ameiva undulata stuarti Smith, Proc. Biol. Soc. 
Washington, 53:55-56, 1940 [Holotype: 
USNM 108601. Type Locality: Palenque, 
Chiapas, Mexico. Collector: H. M. Smith]. 
Stuart, Proc. Biol. Soc. Washington, 55: 
146, 1942. Smith and Laufe, Univ. Kansas 
Sci. Bull., 31(2):50-51, Fig. lb, Pi. lb, 
1946. Smith and Tavlor, U. S. Natl. Mus. 
Bull., 199:173, 1950. 

Ameiva undulata amphigramma Smith and 
Laufe, Trans. Kansas Acad. Sci., 48(3): 
338-344, 1945 [Holotype: FMNH 100020. 
Type Locality: San Andres Tuxtla, Vera- 
cruz, Mexico. Collector: H. M. Smith]. 
Smith and Laufe, Univ. Kansas Sci. Bull., 



31(2):43-47, PI. Ic-Id, 1946. Smith and 
Tavlor, U. S. Natl. Mus. Bull., 199-172, 
19.50. 

Ameiva undulata gaigeae Smith and Laufe, 
Univ. Kansas Sci. Bull., 31(2):37-39, Fig. 
Ic, Pi. lie, 1946 [Holotype: FMNH 
100030. Type Locality: Progreso, Yucu- 
tan, Mexico. Collector: H. M. Smith]. 
Smith and Taylor, U. S. Natl. Mus. Bull., 
199:172, 1950. Stuart, Misc. Publ. Mus. 
Zool., Univ. Michigan, 122:77, 1963. 

Ameiva undulata podarga Smith and Laufe, 
Univ. Kansas Sci. Bull., 31(2):40-43, Fig. 
Id, 2a, 1946 [Holotype: FMNH 100050. 
Type Locality: 7 mi. W Victoria, Tamau- 
lipas, Mexico. Collectors: H. M. Smith 
and D. H. Dimkle]. Smith and Taylor, 
U. S. Natl. Mus. Bull., 199:172, 1950. 

Ameiva imdidata dextra Smith and Laufe, 
Univ. Kansas Sci. Bull., 31(2) :54-,59, Fig. 
2c, 1946 [Holotype: FMNH 100080. Type 
Locality: nr. Rincon, Guerrero, Mexico. 
Collectors: E. H. Taylor and H. M. 
Smith]. Smith and Taylor, U. S. Natl. 
Mus. Bull., 199:173, 1950. 

Ameiva undulata sinistra Smith and Laufe, 
Univ. Kansas Sci. Bull., 31(2):59-62, 1946 
[Holotype: FMNH 100054. Type Locali- 
ty: Manzanillo, Colima, Mexico. Collec- 
tor: H. M. Smith]. Smith and Taylor, 
U. S. Natl. Mus. Bull., 199:174, 19.50. 

Ameiva festiva miadis Loveridge and Barbour, 
Bull. Mus. Comp. Zool., 69(7) :141, 1929 
[Holotype: MCZ 26970. Type Locality: 
Isla del Maiz Grande, Depto. Zelaya, Nica- 
ragua. Collector: James L. Peters]. 

Ameiva undulata miadis — Dunn, Proc. Acad. 
Nat. Sci. Philadelphia, 92:115, 1940. Villa, 
Rev. Biol. Trop., 15(1):119, 1968. Echter- 
nacht, Breviora, 354:5, 1970. 

Distribution. — The most widely clis- 
tiibuted of the Middle American Amei- 
va (Fig. 17), Ameiva umJidata occm-s 
from near Las Varas, Nayarit, Mexico, 
to the vicinity of Quepos, Puntarenas 
Prov., Costa Rica, on the Pacific slopes. 
On the Atlantic slopes, the species 
ranges from Rancho Santa Ana (12.8 
km SE Podilla), Tamaulipas, Mexico, to 
Puerto Lempira, Cabo Gracias a Dios, 
Honduras, and on more westerly Atlan- 
tic slopes of Nicaragua from Corozo, 
Depto. Nuevo Segovia to Tuli Creek, 
Depto. Rio San Juan. Four specimens 
(ANSP 1543S-41) from Huaunta, Depto. 
Zelaya, Nicaragua, seemingly document 
the presence of Ameiva undulata on the 
Caribbean coast of that country (see 



42 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



24^ 



2 



8^ 



5° 



2' 



9' 





£>. 



-24^ 



21 



27-.a^^S^° •34, ,' W 







200 400 



KILOMETERS 



105^ 



99^ 




Fig. 17. Map showing locality records of Ameiva undulata. To avoid crowding of symbols, some 
localities have been omitted. Samples used in statistical analyses were ( 1 ) Gomez Farias and vicin- 
ity, Tamanlipas, Mexico, (2) Tamazunchale and vicinity, San Luis Potosi, Mexico, (3) Tierra 
Colorado, Veracruz, Mexico, (4) Cuautlapam, Veracruz, Mexico, (5) San Andres Tuxtla and 
vicinity, Veracruz, Mexico, (6) Vicinity of Jesus Carran?a, Veracruz, Mexico, (7) Frontera and 

(8) Tres Brazos and Balchacaj, Campeche, Me.xico, (9) Dzilbalchen, 

Piste, Yucatan, Mexico, (11) El Cayo and vicinity, Cayo Dist., British 

and Tikal, Depto. El Peten, Guatemala, (13) Piedras Negras, Depto. 

Canihor, Depto. Alta Verapaz, Guatemala, (15) El Rancho and Finca 



vicinity. Tabasco, Mexico, 
Campeche, Mexico, ( 10 ) 
Honduras, ( 12 ) Uaxactiin 
El Peten, Guatemala, (14) 



Bucural, Depto. Progreso, Guatemala, (16) Copan, Depto. Colon, Honduras, (17) La Ceiba, 
Depto. Atlantida, Honduras, (18) Coyoles Central, Depto. Yoro, Honduras, (19) Trujillo, Depto. 
Colon, Honduras, (20) Catacamas, Depto. Olancho, Honduras, (21 ) Isla del Maiz Grande, Depto. 
Zelaya, Nicaragua, (22) Manzanillo-Pacific Coast Localities, Colima, Mexico, (23) Colima and 
vicinity, Colima, Mexico, (24) Acapulco and vicinity, Guerrero, Mexico, (25) Chilpancingo and 
vicinity, Guerrero, Mexico, (26) Chacalapa and vicinity, Oaxaca, Me.xico, (27) Tres Cruces, 
Oaxaca, Mexico, (28) Tolocito, Oaxaca, Mexico, (29) Tapanatepec, Oaxaca, Mexico, (30) Vicinity 
of Cintalapa, Chiapas, Mexico, (31) Las Tazas, Chiapas, Mexico (51 km E Altamirano), (33) 
Esquintla, Chiapas, Mexico, (34) Sabana de San Quintin, Chiapas, Mexico, (35) Panajachel, 
Depto. Solola, Guatemala, (36) Finca La Trinidad, Depto. Jutiapa, Guatemala, (37) JuHapa 
and Finca Mongoy, Depto. Jutiapa, Guatemala, (38) Vicinity of La Libertad, Depto. La Liber- 
tad, El Salvador, (39) Chinandega, Depto. Chinandega, Nicaragua, (40) Matagalpa, Depto. 
Malagalpa, Nicaragua, (41) Hacienda La Norma and vicinity, Guanacaste Prov., Costa Rica, (42) 
Canas, Guanacaste Prov., Costa Rica, (43) Tilaran, Guanacaste Prov., Costa Rica, (44) El Zamo- 
rano and vicinity, Depto. Francisco-Morazan, Hondmas, (46) Isla Mujeres, Quintana Roo, Mex- 
ico, (48) Apatzingan, Michoacan, Mexico. Open symbols indicate literature records of specimen-S 

not examined for the present study. 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



43 




Fig. 18. Head and preanal scutellation of 
Ameiva nndulata (KU 94112; Sabana San 
Quintin, Chiapas, Mexico): (A) Lateral view 
of head, (B) Dorsal view of head, (C) Ventral 
view of head, X 1.1- (D) Preanal region, 
X 1.5. 



Remarks). The species is known from 
Isla Mujeres, east of Quintana Roo, 
Mexico, and from the Islas del Maiz 
east of Nicaragua. Gene flow between 
populations on the Atlantic and Pacific 
slopes occurs across the Isthmus of Te- 
huantepec, in the lowlands of eastern 
Guatemala, and in Honduras and Nica- 
ragua south of the highlands that mark 
Nuclear Central America. Ameiva tin- 
chiJata is generally found at elevations 
below 1500 m, although there are ex- 
ceptions (e.g., at Panajachel, Depto. 
Solola, Guatemala). 

Diagnosis. — Ameiva umJulata can be 
distinguished from other Middle Ameri- 
can Ameiva by the following combina- 
tion of characters: Gentral gular scales 
slightly to greatly enlarged, regular or 
irregular in arrangement; gradual re- 
duction in size of scales radiating out- 



ward from central gular scales; prefron- 
tal scales in contact with postnasal 
scales; three parietal scales (four if me- 
dian parietal divided); a transverse row 
of abruptly enlarged mesoptychial 
scales; usually eight rows of ventral 
scutes; a relatively broad middorsal 
stripe bounded by narrow light stripes 
which may be lost in adults; no narrow 
vertebral stripe; lateral pattern empha- 
sizes longitudinal stripes or vertical bars 
in adults, never spots. 

Description. — Maximum observed 
SVL 129 mm for males, 111 mm for 
females; supralabials 12-18 (14.1) for 
males, 12-18 (14.1) for females; infra- 
labials 7-14 (10.2); supraoculars 6-10 
(6.4); COF values 2-6 (2.2); GOP 
values 2-14 (10.6); SO-SG values 2-10 
(3.8); loreals 2-8 (4.0); nostril in pre- 



nasal-postnasal suture; 
contact with postnasal; 



prefrontal in 
median gular 




Fig. 19. Head and preanal scntellation of 
Ameiva unchdata (KU 26963; 25 km SE Jesus 
Carranza, Veracruz, Mexico): (A) Lateral 
view of head, (B) Dorsal view of head, (C) 
Ventral view of head, X 1.1. (D) Preanal 
region, X 1-5. 



44 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 




Fig. 20. Preanal Region of Ameiva undulata 
to illustrate two of the pattern types: (A) 
KU 87417; East of San Andreas de la Cruz, 
Guerrero, Mexico; Preanal scales in a single 
row except for paired terminal scales, X 2.7. 
(B) KU 38232; Chacalapa, Oaxaca, Mexico; 
Preanal scales in a single row including single 
terminal preanal, X 1.5. 

scales variable in size and orientation; 
GAB 108-202 (149.5); PV 31-67 (47.4) 
for males, 33-65 (46.4) for females; 
PV/GAB 0.21-0.40 (0.32) for males, 
0.23-0.43 (0.31) for females; GOR 186- 
315 (246.0); GAB/GOR 0.44-0.77 
(0.61) for males, 0.49-0.74 (0.60) for 
females; SAT 18-26 (22.3) for males, 
18-27 (22.2) for females; total preanal 
scales 4-13 (7.6) for males, 5-13 (7.5) 
for females; terminal preanal scales 1-3 
(2.0); one (usually) or two rows of en- 
larged prebrachials; postbrachials en- 
larged, in one row; preantebrachials in 
two rows, one row or two rows prox- 
imally and one row distally; prefemoral 
scales enlarged, three to five rows prox- 
imally reducing to two (usually) or 
three rows distally; infratibials enlarged, 
in two rows, femoral pores 29-48 (37.4) 
for males, 22-45 (35.0) for females; sub- 
digital lamellae 24-38 (30.2); longitu- 
dinal rows of ventral scutes 26-34 
(30.1); transverse rows of ventral scutes 
8-12 (8.2). For details of scutellation 
see figures 18-20. 

Color and Pattern in Alcohol. — 
Throughout its broad range, Ameiva 
undulata shows geographic variation in 
color and pattern unmatched by other 
Middle American species of Ameiva 
and ontogenetic variation, especially of 
males, greater than all, except possibly 
A. ameiva. The following description 
of color and pattern in alcohol, as well 
as the description of color and pattern 



in life, is based on specimens of undu- 
lata from the vicinities of La Ceiba and 
Trujillo along the north coast of Hon- 
duras (Localities 17 and 19, Fig. 21). 

Juvenile Males: Top of head brown; 
side of head black above, grading 
through brown to gray on supralabials; 
a white patch beneath and slightly be- 
hind eye; ear surrounded by white 
markings; middorsal field yellowish- 
brown or greenish-brown with black 
flecking, bordered laterally by fine white 
cream-colored dorsolateral stripes which 
originate above eye and extend well 
onto tail; dorsolateral dark stripe ex- 
tending from behind eye to base of tail, 
velvet black with no light bars or 
blotches, bounded below by an inter- 
rupted white ventrolateral light line 
originating behind the ear and extend- 
ing to the groin, thence as an uninter- 
rupted white line well onto tail, ventro- 
lateral dark field gray-brown; dorsal 
surfaces of limbs mottled black and 
brown, hind limbs with white spots 
above; white postfemoral stripe joining 
caudal extension of ventrolateral light 
stripe; tail brown with black flecks 
above; ventral surfaces immaculate or 
blue-gray. 

Through ontogeny (Fig. 21) the 
dorsolateral light stripe disappears or is 
retained only on the anterior half of the 
body. The ventrolateral light stripe is 
usually lost entirely. Light blue vertical 
bars appear on the flanks, extending 
from the border of the middorsal field 
or near it to or nearly to the enlarged 
ventral scutes. Shorter bars and blotches 
may appear between the primary bars. 
The light spots around the ear are lost 
and the venter may become light blue 



Fig. 21. Ontogenetic change in pattern of 
Avieiva undulata from northern, coastal Hon- 
duras. Males: (A) KU 101247, SVL 71 mm; 
(B) KU 101238, 92 mm; (C) KU 101237, 
110 mm. Females: (D) KU 101242, 39 mm; 
(E) JRM 2318, 88 mm; (F) JRM 2554, 109 
mm. Small males resemble small females as 
exemplified by KU 101242 (D above). 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



45 




46 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



overall except for black patches in the 
lateral-most scales and on the enlarged 
plates on the anterior surface of the 
femur. Traces of red pigment are visi- 
ble in the gular region of some large 
males. 

Females: Juvenile females are simi- 
lar to juvenile males. Through ontog- 
eny, females generally retain the juve- 
nile pattern, except that the dorsolateral 
light stripe may be lost on the pos- 
terior one-half of the body and regu- 
larly spaced light brown blotches ap- 
pear in the dorsolateral dark field (Fig. 
21). 

Color and Pattern in Life. — The fol- 
lowing is based on field notes and color 
photographs of Ameiva tindulata col- 
lected during July, 1967, near La Ceiba 
and Trujillo on the north coast of Hon- 
duras. 

Adult Males: Top of head gray- 
brown to light brown; middorsal field 
gray-green on neck becoming bright 
leaf-green on shoulder and extending 
one-half to two-thirds length of trunk, 
then becoming golden-brown on pos- 
terior part of body; entire middorsal 
field irridescent in sunlight; dorsal sur- 
face of tail gray-brown with black fleck- 
ing; middorsal field from shoulder to 
tail lightly to moderately blotched with 
black which sometimes appear paired; 
remnant of dorsolateral light stripe 
cream, if present, and only visible on 
anterior two-thirds of body; lateral pat- 
tern of seven to ten vertical blue-white 
or greenish-white bars which may touch 
middorsal field but which usually are 
separated from it and which taper so 
that they are wider than or equal to the 
interspaces dorsally but much narrower 
than interspaces ventrally; short blue 
bars extending upward between the 
prominent lateral bars, sometimes con- 
necting with them; lateral ground color 
black above (in area of dorsolateral 
dark field of juveniles), reddish-brown 
below. Some individuals retain a rem- 
nant of the cream-colored ventrolateral 
light stripe so that the dorsolateral dark 



field may be defined. Dorsal surfaces of 
limbs brown with indistinct black mot- 
tling; ventral coloration of limbs, body 
and tail blue-white with black flecking 
and blotches, especially on the hind 
limbs and laterally on the body; chin, 
gular region and neck range in color 
from white to brick red with no indica- 
tion of yellow pigmentation. 

Adult Females: Dorsal pattern simi- 
lar to that of males except that cream- 
colored dorsolateral light stripes usually 
are retained; dorsal coloration similar to 
that of males but several shades darker 
overall; ventrolateral light stripe cream, 
continuous, broken or broken anteriorly, 
entire posteriorly; dorsolateral dark field 
black with brown or cream blotches, 
each wider than or equal to width of 
interspaces; ventrolateral field brown or 
gray-brown with darker mottling; dor- 
sal surfaces of limbs as for males; ven- 
tral surfaces immaculate or with slight 
bluish tint laterally and on hind legs. 
Females lack the bright vertical bars on 
the flanks, never show reddish pigment 
on the ventral surfaces of the head and 
neck and never develop blue ventral 
coloration as markedly as do males. 

Geographic Variation. — The follow- 
ing samples were removed during STP 
analysis of sexually dimorphic charac- 
ters because of inadequate sample sizes: 
For characters of males, samples 3, 9 
and 48; for characters of females, sam- 
ples 7, 9, 23, 25, 26, 34, 38, 46 and 48. 
In addition, sample 3 was removed in 
the analysis of number of loreals of 
females and both 3 and 22 were re- 
moved in the analysis of SO-SC values 
of both sexes, again for reasons of inade- 
quate sample size. 

The most obvious patterns of geo- 
graphic variation for meristic characters 
of Ameiva undulata are clinal trends 
along the northeast and northwest coasts 
of Mexico and along the Pacific coast of 
Middle America from the Isthmus of 
Tehuantepec to Nicaragua and Costa 
Rica ( Fig. 22 ) . In some cases one sam- 
ple along such a cline is "out of phase," 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



47 



interrupting what otherwise is a rather 
smooth trend. Such samples are indi- 
cated in parentheses in the following 
account. 

Those characters showing an in- 
crease in sample means from Tamaulipas 
to the Isthmus of Tehuantepec include 
number of supraoculars (2), number of 
paravertebral granules of males, num- 
ber of paravertebrals of females (3), 
total preanal scales of males, total pre- 
and scales of females (4) and number 
of femoral pores of males. Shorter 
clines in the same direction, but involv- 
ing only samples 1 through 3 or 4 are 
apparent for PV/GAB of males and 
SAT of males (the size of sample 3 is 
too small for inclusion in the analysis 
for both characters), as well as number 
of femoral pores of females. The cline 
for total preanal scales of males and of 
females extends southward across the 
isthmus and thence eastward along the 
Pacific coast as far as locality 33 for 
males and locality 36 for females. Two 
characters, COF values and number 
loreals, show clinal increases from north 
to south across the isthmus and along 
the coast to varying degrees. For both 
characters, lowest mean values are for 
sample 5. COF values increase through 
samples 28 and 29 and through samples 
6, 30 and 33 to 36. The number of 
loreals increases through samples 28, 27 
and 26 and through samples 28, 29 and 
30. Mean numbers of supraoculars 
show the same trans-isthmian pattern 
as COF values, but in the opposite di- 
rection and with the lowest value at 
sample 35, not 36. CAB increases from 
locality 5 through 28 and 29 or 6 and 
30 to 33 and 36. 

Decrease in mean values from Ta- 
maulipas to the Isthmus of Tehuantepec 
is notable for the number of supralab- 
ials of females. Shorter clines in the 
same direction but involving only sam- 
ples 1 through 3 or 4 are apparent for 
CAB and CAB /COR of males (sample 
3 is too small for inclusion in the analy- 
sis of CAB /COR of males) and number 




Fig. 22. Major patterns of geographic varia- 
tion in Ameiva ttndulata. Arrows represent 
geographical trends involving several charac- 
ters. Samples on one side of the heavy, dashed 
line differ from adjacent samples on the other 
side of the line in a statistically significant 
manner for five or more characters but do not 
differ in such a way from adjacent samples on 
the same side of tlie line. See text for further 
elal)oration. 



of subdigital lamellae. Both COP and 
SO-SC values increase from south to 
north across the isthmus and a west to 
east and northeast increase is noted for 
SO-SC values. 

Northwest to southeast clines are 
apparent along the south coast of Mex- 
ico also. Characters showing increased 
sample means from Colima to the Isth- 
mus of Tehuantepec or at least to the 
vicinity of Chacalapa, Oaxaca (Sample 
26) include number of paravertebral 
granules of males and of females, PV/ 
CAB of males and of females, total pre- 
anal scales of females and SAT of males 
(inadequate sample size precludes in- 
clusion of sample 48 in analysis of these 
characters). Although the clinal trend 
terminates at the isthmus for the other 
characters mentioned, that for total pre- 
anals of females continues along the 
Pacific coast to locality 36 in Cuatemala. 
A short trend in the same direction is 
noted for number of transverse rows of 
ventral scutes from localities 22 and 23 
through 48 to 24 and 25. 

Trends along the same coast, but 
showing a decrease in sample means 
from Colima to the Isthmus of Tehuan- 



48 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



tepee are noted for number of siipra- 
labials of males (sample 48 excluded 
from the analysis) and females (sam- 
ples 26 and 48 excluded), number of 
infralabials, GAB/GOR of females, 
number of tenninal preanals and num- 
ber of subdigital lamellae. These trends 
generally involve only coastal samples 
(samples 22, 24, 26, 27 and 29), but in- 
clude inland samples (samples 23, 25 
and 48) for supralabials of males and 
total number of infralabials. 

A number of characters show clinal 
tendencies for increase or decrease in 
sample means between Nicaragua or 
Costa Rica and localities 33, 37 and 
especially 36 to the northwest. There is 
a marked tendency for clines to either 
begin or end in the area of southeastern 
Mexico and southern Guatemala with 
no clines running through to the south- 
east or northwest along the coast. Num- 
bers of supraoculars show an increase 
from sample 40 (.x = 6.13) in Nicara- 
gua to sample 36 (x = 9.97) in Guate- 
mala, at which point there is an abrupt 
decrease in sample mean and a clinal 
increase from sample 33 (x = 6.28) to 
sample 5 ( x = 6.70 ) on the Gulf side of 
the Isthmus of Tehuantepec in Mexico. 
Samples 34 (Ghiapas) and 35 (Guate- 
mala) are significantly different from 
the nearest samples to the east in Guate- 
mala for this character, but sample 36 
(Guatemala) is not significantly differ- 
ent from sample 33 (Chiapas). A simi- 
lar trend for increase is noted between 
sample 39 (x = 2.07) in Nicaragua and 
sample 37 (x = 2.29) in Guatemala for 
COF values. Sample 36 (x = 3.77), 
located just west of sample 37, is signifi- 
cantly different from it and is the end of 
a cline beginning at sample 5 (x = 
2.13) and which progresses across the 
Isthmus of Tehuantepec through sam- 
ples 28 and 29 or samples 6, 30 and 33. 
Samples 33 and 36 have very high mean 
values for COF and are statistically iso- 
lated from samples to the north. Fem- 
oral pore means for both males and 
females also show increases from sam- 



ple 40 to sample 37, but the trend may 
be traced from samples 41, 42 and 43 in 
Costa Rica as well. For this character 
in females the cline continues to sample 
15 and may also be traced from the 
Costa Rican samples or sample 40 in 
Nicaragua through samples 39 and 44 
to sample 16 in western Honduras. The 
number of supralabials of males in- 
creases from sample 37 north and west- 
ward to sample 29, and GAB/GOR for 
females increases from sample 37 to 
sample 33 and from sample 15 to sam- 
ple 33, both trends running to the west 
or northwest. Mean numbers of sub- 
digital lamellae increase from samples 
20 and 44 through samples 38 and 37 to 
sample 36 on the coastal plain of Guate- 
mala and to sample 33 by way of sam- 
ples 15 and 35. Total number of pre- 
anal scales of females follow the same 
trend. These two characters are the 
only ones involved in clines running 
through the area of samples 36 and 37. 
Mean numbers of supralabials of fe- 
males increase from west to east, be- 
ginning at sample 30 and terminating at 
samples 14 and 37 by way of samples 
33, 35 and 15. Sample 36 (x = 13.5), 
with a mean for this character lower 
than that of sample 30 (x ^ 13.6) is not 
part of the trend. 

Mean numbers of paravertebral 
granules of males and of females and 
PV/GAB of females show trends for in- 
crease from sample 40 in Nicaragua or 
sample 20 in Honduras southwestward 
and along the Pacific coast to sample 36 
in Guatemala. With respect to number 
of paravertebrals of females, sample 36 
is significantly different from sample 37. 
For both number of paravertebrals of 
females and PV/GAB of females, the 
cline may be traced to either sample 36 
or sample 15 in Guatemala. 

Mean SO-SC values and mean num- 
ber of loreals show increased values 
from north to south along a line be- 
tween sample 9 on the Yucatan Penin- 
sula and samples 37 and 38 in south- 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



49 



eastern Guatemala and El Salvador, 
respectively. 

There seems to be a trend toward 
increase in mean numbers of paraverte- 
bral granules of females, GAB/GOR of 
females and total number of preanal 
scales of males from sample 8 in south- 
western Campeche through sample 10 
in Yucatan to sample 46 on Isla Mujeres. 
There is a similar trend beginning at 
sample 7 in Tabasco and terminating at 
sample 10 for mean numbers of femoral 
pores of males. For all of these charac- 
ters, however, sample 9 in central Cam- 
peche ( Dzilbalchen ) is too small for 
meaningful comparison (n = 3) where 
females are concerned and such trends 
might owe their existences to lack of 
data from this locality. A clinal de- 
crease from sample 8 through sample 9 
and 10 to sample 46 is apparent for 
mean number of rows of transverse ven- 
tral scutes. An increase along the same 
line is noted for mean number of 
loreals. 

Most samples differ from adjacent 
samples in a statistically significant 
manner for at least one character. For 
purposes of this discussion, a character 
judged sexually dimorphic would count 
as two characters because separate con- 
sideration is given to males and females. 
If, for example, two samples were sig- 
nificantly different with respect to num- 
ber of femoral pores of males and fe- 
males, they would be said to differ sig- 
nificantly for two characters, not one. 
Viewing all characters at once gives one 
the impression that each sample is 
more-or-less statistically isolated from 
its neighbors. Although this may be 
true, it renders analysis and formation 
of meaningful generalizations extremely 
difficult. However, if only pairs of sam- 
ples differing statistically by (arbi- 
trarily) five or more characters are 
examined, some interesting patterns 
emerge. These are discussed below. It 
must be emphasized that comparisons 
are, in most cases, being made only be- 
tween adjacent samples. Two samples 



adjacent to one another in Honduras 
may be statistically different from one 
another on the basis of one or more 
characters, but neither may differ from 
a sample in, for example, northern 
Mexico. Of course, the opposite is also 
possible. 

The population of Ameiva iindulata 
on Isla del Maiz Grande (sample 21) 
differs from samples 42, 43 and 44 for 
five characters each, from samples 39 
and 40 six characters each and from 
sample 20 for ten characters. With the 
exceptions of samples 40 and 41, which 
differ from each other for five charac- 
ters, none of these mainland samples 
differ from adjacent samples among 
those mentioned by more than four 
characters. This strongly emphasizes 
the divergence of the Ameiva on Isla 
del Maiz Grande from the nearest main- 
land populations, or at least from the 
available samples of those populations, 
and is indicated by the heavy, dashed 
line in figure 22. Although sample 20 
differs from sample 21 for twice or more 
the number of characters separating 
sample 21 from other mainland samples, 
there is still little basis for concluding 
that A. undulata on the Islas del Maiz 
were derived from elsewhere because 
the Honduranian population from which 
sample 20 was drawn may have been 
modified itself through introgression of 
characteristics of populations to the 
north and west (samples 17, 18 and 19). 

The latter three samples represent a 
population quite distinct from neigh- 
boring samples on the basis that none 
of the three differ among themselves by 
more than one character ( 17 and 18 dif- 
fer from 19 for SO-SG values; other 
characters show non-significant differ- 
ences among sample means), but all 
differ from other nearby samples by 
more than five characters (see Fig. 22). 
Samples 16, 20 and 44, which encircle 
samples 17, 18 and 19, do not differ 
among themselves, but all show con- 
siderable divergence from samples 17, 
18 and 19 for meristic and, as will be 



50 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



discussed later, color and pattern char- 
acters. Samples 17, 18 and 19 differ 
from sample 11 (El Cayo and vicinity, 
British Honduras) for only three, one 
and one characters, respectively. Sam- 
ple 16, on the other hand, differs from 
sample 11 for four characters, none of 
which are among those few which dis- 
tinguish sample 11 from samples 17, 18 
and 19. 

Samples 9-16, 31, 34, 35, 37, 38 and 
46 all show statistically significant dif- 
ferences involving fewer than five char- 
acters when compared among them- 
selves with respect to adjacent samples. 
However, sample 8 differs from samples 
12, 13, 31, 33 and 34 for ten, ten, eight, 
ten and seven characters, respectively. 
In addition to sample 8, sample 31 dif- 
fers from samples 6, 7, 30, 33 and 35 for 
six, seven, nine, ten and nine characters 
respectively. Sample 33, in addition to 
differing from samples 8 and 31 by 
more than five characters, differs from 
samples 34 and 35 by six and seven 
characters, respectively. Sample 36 dif- 
fers from samples 15, 35 and 37 for 
seven, eight and seven characters, re- 
spectively. Samples 8, 7, 6, 30, 33 and 
36 constitute a series of localities be- 
ginning in the Tres Brazos-Balchacaj 
area of Campeche, progressing west 
along the Gulf coast and thence south 
across the Isthmus of Tehuantepec and 
eastward along the Pacific slopes to the 
vicinity of Finca La Trinidad, Depto. 
Jutiapa, Guatemala. These samples, as 
indicated above, all differ from adjacent 
more inland samples to the south, east 
or north by five or more characters. If 
a line is drawn to emphasize the diver- 
gence of these "peripheral" samples 
from the other, more inland, samples 
(Fig. 22), none of the samples falling 
within the curvature of the line differ 
from adjacent samples by as many as 
five characters with the exception of 
sample 15, which differs from sample 35 
for six characters. 

Sample localities from the northern 
Gulf region of Mexico (samples 1-5) are 



isolated from sample localities along the 
southwest Pacific coast (samples 22-27, 
48) by the Mexican Plateau. In gen- 
eral, however, samples on the one coast 
differ for five or more characters only 
from the most distant samples on the 
other coast. For example, sample 1 
(Tamaulipas) differs from sample 27 
(south-central Oaxaca) for seven char- 
acters, but differs from sample 22 (Co- 
lima) for only three characters. Sample 
5 ( San Andres Tuxtla and vicinity, 
Veracruz) differs from sample 27 for 
onlv four characters, but differs from 
sample 22 for nine. Samples 1-5 and 
22-27, 48 do not differ from adjacent 
samples on the same coast by more than 
two characters, with the exception of 
samples 25 and 26, which differ for four 
characters. 

Sample 27 (Tres Cruces, Oaxaca), 
however, seems to be strongly diver- 
gent from adjacent samples to the east. 
Sample 27 differs from samples 6, 28 
and 29 for ten, six and eight characters, 
respectively. This, coupled with the 
isolation afforded by the Mexican Pla- 
teau, is the basis for separating off the 
samples from the Pacific slopes of Mex- 
ico northwest of the Isthmus of Tehuan- 
tepec as shown in figure 22. 

The groupings of samples set apart 
according to the method just described 
fits well with the general pattern of 
clinal trends discussed earlier. There is 
also a generally good fit by these group- 
ings with the placement of highland 
regions with elevations above 1500 
meters, an elevation above which Amei- 
va rarely occur in Middle America. 

Using the 14 samples (8, 15-21, 27, 
31, 33, 35, 37 and 44) which were found 
to differ from adjacent samples by five 
or more characters, it is possible to ar- 
rive at some idea of which characters 
are most discriminatory in distinguish- 
ing adjacent samples. The 14 samples 
were compared with adjacent samples 
to give 68 different comparisons. It was 
found that in 53 per cent of these com- 
parisons (36 out of the possible 68), 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



51 



mean GAB differed in a statistically 
significant manner. GAB is apparently 
the primary meristic indicator of sam- 
ple differences, followed by GOR (43 
per cent of the comparisons), SO-SC 
values (31%), number of subdigital 
lamellae (31%) and COP values (29%). 
All other characters were involved in 
26 per cent or less of the comparisons 
each. Two characters, number of su- 
pralabials of males and number of trans- 
verse rows of ventral scutes, were found 
not to distinguish any of the 68 pairs of 
samples compared. All five of the char- 
acters which seem most important in 
this respect do not appear together in 
distinguishing two adjacent samples, 
but they do appear in groups of 2, 3 or 
4 in various comparisons. GAB and 
GOR, as might be expected, occur to- 
gether most often in distinguishing pairs 
of adjacent samples, appearing in 26 of 
the 68 comparisons (38%). GAB and 
number of subdigital lamellae occur to- 
gether in distinguishing 15 pairs among 
the 68 (22%), whereas GAB and SO-SC 
values, and GAB and COP values both 
occur together in 14 of 68 comparisons 
(20%). Other combinations of the five 
characters taken two at a time occur 
less frequently. 

The condition of the preanal scales 
of Ameiva tindtilata can be analyzed in 
a number of ways. Two of these, the 
total number of preanals and the num- 
ber of terminal preanal scales have 
been defined and discussed with respect 
to STP analysis. These characters, along 
with a third which combines the two, 
warrant further discussion. 

For STP analysis, the number of 
terminal preanal scales was coded 1, 2 
or 3, indicating one, two or three scales 
in the terminal position. An inherent 
weakness in this system is that a sample 
mean or range gives no indication of 
the actual frequency distribution of 
single, double and triple terminal pre- 
anals in the sample. A sample of two 
individuals, one having one terminal 
preanal, the other having three, would 



have the same mean number of terminal 
preanals as another sample of two in- 
dividuals where each animal had two 
terminal preanal scales. This is an over- 
simplification complicated by larger 
sample sizes. Even statistics, such as 
standard deviation, are of little help in 
visualizing the frequency distribution. 
For this reason, the number of terminal 
preanal scales was coded so that three 
terminal preanal scales received the 
same value as one. This system has 
some validity in that only scales in the 
midline are being compared. Individ- 
uals having two terminal preanals con- 
tinue to receive a code value of two. If 
this is done, most samples still have a 
mean number of tenninal preanals near 
or approaching 2.00, but certain sam- 
ples have much lower values. The low- 
est of these are samples 1 (x = 1-67), 
8 (1.60), 17 (1.67), 22 (1.42), 23 
(1.61), 25 (1.76), 26 (1.66), 27 (1.33) 
and 48 (1.78). Means of other samples 
ranged from 1.80 (sample 9) to 2.00 
(many samples). It will be noted that 
most of the low sample means are en- 
countered in samples 22-27 and 48 be- 
tween Colima and Tres Cruces, Oaxaca, 
in Mexico. Sample 24, consisting of 14 
individuals each with two terminal pre- 
anal scales, does not fit the pattern. 

Coupled with the number of termi- 
nal preanal scales is the pattern of these 
scales. They may be (A) mostly 
(>50%) single (Fig. 20B), (B) mostly 
paired, (C) single except the terminal 
scales paired (Fig. 20A), or (D) mixed. 
The latter category includes individuals 
not fitting into any other category if the 
single or paired condition is based on 
three or more scales or pairs of scales in 
a row. Most individuals of Ameiva tin- 
dtilata fall into class C (71.6%; n r= 
1003), whereas fewer are referable to 
classes A (18.2%), B (4.5%) and D 
(5.7%). Two series of samples, how- 
ever, show marked deviation from this 
distribution. If samples 22-27 and 48, 
from the Pacific slopes of Mexico north 
and west of the Isthmus of Tehuan- 



52 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



tepee, are eombined, there are about 
equal numbers of individuals referable 
to classes A (33.9%), B (27.3%) and D 
(30.6%), but only 8.3 percent are refer- 
able to class C (n = 121). Individual 
lizards in samples 41-43 from Costa 
Rica are distributed as follows: class 
A (44.2%); class B (3.9%); class C 
(35.1%); class D (16.9%) (n = 77). 
On the basis of this character alone 
samples 41-43 differ markedly from 
samples to the north (samples 39-40 in 
Nicaragua), but the divergence of sam- 
ples 22-27 and 48 parallels that noted 
for number of terminal preanal scales 
and appears to reinforce the evidence 
for a break in gene flow between popu- 
lations of iindiilata east and west of a 
line drawn between sample 27 and 
samples 28 and 29. If samples 21-27, 
41-43 and 48 are removed, the distribu- 
tion of individuals in the remaining 
samples becomes: class A (13.4%); 
class B (1.1%); class C (84.6%); class 
D (0.9%) (n = 805). 

Another character not analyzed by 
STP, but warranting further mention is 
the pattern of the median gular scales. 
Median gular scales may be either ir- 
regular in arrangement or oriented 
longitudinally in a more-or-less regular 
row. If irregular, they are usually at 
least slightly" enlarged (39.0%; 404 of 
1026 individuals), although a few in- 
dividuals have midgular scales no larger 
than surrounding scales (0.4%). If 
longitudinally oriented and enlarged, 
they may be paired (2.1%), in a row of 
single scales (34.8%) or mixed paired 
and single (23.1%; at least three scales 
or pairs of scales in a row were deemed 
necessary to classify a lizard as having 
longitudinally oriented scales). Sam- 
ples 3-8, 15, 22-30, 33, and 35-36 are 
made up of individuals more than 70 
percent of which have enlarged, longi- 
tudinally oriented gulars. Samples 5, 7, 
8 and 28 contain only such individuals. 
These sample localities are distributed 
along the Gulf coast of Mexico east and 
west of the Isthmus of Tehuantepec 



from northwestern Veracruz to the Tres 
Brazos-Balchacaj region of Campeche, 
across the Isthmus and along the Pacific 
coast from Colima to southeastern 
Guatemala (Fig. 17). Samples 9-14, 17- 
19, 31 and 34 from the Yucatan Penin- 
sula, eastern Mexico, northern Guate- 
mala and northern Honduras are com- 
posed of individuals more than 70 per- 
cent of which have enlarged, irregularly 
arranged gulars. Samples 13 and 34 
consist of only such individuals. 

These patterns are generally re- 
flected in the lines between adjacent 
samples depicting significant differences 
involving five or more characters (Fig. 
22). Gular pattern tends to place sam- 
ples 17-19 from northern Honduras with 
samples from northern Guatemala and 
Yucatan Peninsula. The line separating 
samples 6-8, 30 and 33 from samples 12- 
13, 31 and 34 is represented, but sam- 
ples 15 and 35 are not differentiated 
with respect to gular scale pattern and 
these samples do not differ from sample 
36. Samples 15, 35 and 36 differ from 
one another for at least five of the char- 
acters analyzed by STP. With respect 
to gular pattern, sample 36 differs 
sharply from sample 37 and most other 
samples to the south and east. Sample 
36 contains only one individual out of 
30 (3.3%) with irregularly arranged 
gulars, whereas sample 37 contains 13 
of 31 (41.9%) individuals with irregular 
gular scales. 



Fig. 23. Selected geographic variants of color 
pattern of adult Ameiva undulata. (A) KU 
101300, male, SVL 126 mm, (B) KU 101301, 
female, 109 mm, both from Isla del Maiz 
Grande, Depto. Zelaya, Nicaragua; (C) KU 
87409, male, 93 mm, from 5 km E Vista Her- 
mosa, Oaxaca, Mexico; (D) UMMZ 41435, 
female, 99 mm, from Cuautlapam, Veracruz, 
Mexico. (E) KU 95645, male, 92 mm, from 
3 km E Mapastepec, Chiapas, Mexico; (F) KU 
95643, female, 82 mm, from 4.5 km E Mapa- 
stepec, Chiapas, Mexico. (G) KU 101260, 
male, 93 mm, (H) KU 101267, female, 90 
mm, both from El Zamorano, Depto. Fran- 
cisco- Morazan, Honduras. 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



53 




54 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



Samples 16 ( 19% with irregularly 
arranged gvilars; n = 16), 40 (20%; n 
= 15) and 2 (73%; n = 15) stand out 
as having unusually high or low num- 
bers of individuals with irregularly 
arranged gular scales relative to adja- 
cent samples. Sample 2 would seem to 
belong with samples from the Yucatan 
Peninsula, northern Guatemala and 
northern Honduras, whereas samples 16 
and 40 are more like samples involved 
in the Isthmus of Tehuantepec "X." 

In samples other than those spe- 
cifically mentioned above, the percent- 
age of irregularly arranged gular scales 
ranged from 33 to 63. 

Discussion of geographic variation 
in color and pattern over the broad 
range of Ameiva undulata is compli- 
cated by the presence of both onto- 
genetic variation (especially among 
males) and sexual dimorphism (Stuart, 
1942). The magnitude of ontogenetic 
change encountered is apparent in fig- 
ure 21, and sexual dimorphism in pat- 
terns of adults from selected popula- 
tions is represented in figure 23. That 
geographic variation is considerable is 
also apparent from figure 23. There are 
two basic pattern types (Stuart, 1942). 
The first of these is characterized by 
an emphasis on longitudinal stripes 
rather than blotches or bars on the 
flanks, by the small amount of sexual 
dimorphism evident, and by its re- 
stricted geographic range. This pattern 
type occurs only in samples 7 and 8 of 
the present study, but it apparently 
grades into other pattern types to the 
west and south. Smith and Laufe ( 1946, 
PI. IB) have illustrated an individual 
exhibiting this color pattern from Teno- 
sique. Tabasco, Mexico. These authors 
cite specimens having this pattern from 
Palenque and San Ricardo, Chiapas; 
Frontera, Teapa and Tenosique, Ta- 
basco; Balchacaj and Tres Brazos, Cam- 
peche. I have examined two specimens 
from San Ricardo, Chiapas (FMNH 
106715, male; UIMNH 26157, female). 
The male has 10 light-colored blotches 



in the dorsolateral dark field, a pattern 
not characteristic of the striped forms 
of undulata. Nine males (FMNH 
106715; UIMNH 8437, 8439, 30229, 
39233-4, 39237-40) from Finca San Bar- 
tola, ca 12 mi SW of Cintalapa and 
near San Ricardo, also show prominent 
bars and/or blotches on the flanks. The 
specimens from Finca San Bartola and 
San Ricardo comprise sample 30 of this 
study; on the basis of both meristic and 
pattern characters, this sample is dis- 
tinct from those to the north. The 
striped pattern type seems to be re- 
stricted to the lowlands of Tabasco, ex- 
treme western Campeche and northern 
Chiapas. The pattern is not unlike that 
of quadrilineata (Fig. 16) of Nicaragua, 
Costa Rica and Panama or chaitzami 
(Fig. 27) of eastern Chiapas, Mexico 
and Guatemala. Some undulata having 
the striped pattern share with chaitzami 
and, to a much lesser extent, quadrili- 
neata, the presence of a secondary dark 
stripe in the middorsal field medial to 
the dorsolateral light stripes. 

The second basic pattern type is 
characterized by a highly variable pat- 
tern of bars and /or blotches on the 
flanks of adult males often accompanied 
by loss of the ventrolateral light stripe 
or its reduction to a series of dashes or 
spots. However, some adult females, as 
well as juveniles of both sexes, retain a 
pattern of longitudinal stripes ( Figs. 
21, 23). Specimens of Ameiva undidata 
included in samples 4, 6 and 28 possess 
this pattern type (Fig. 23 C and D; 
Smith and Laufe, 1946, Plate I C and 
D). The dorsolateral dark field is re- 
tained in both sexes, but the ventro- 
lateral light stripe may be completely or 
partially lost in adult males such that 
the ventral border of the dark field is 
indistinct. The ventrolateral light stripe 
is usually retained in females. The dor- 
solateral light stripe, which is usually 
retained in females may be completely 
lost in males or retained only anteriorly. 
In many specimens the middorsal field 
contains dark patches which lie para- 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



55 



vertebrally iind may be paired. This is 
especially notable in specimens from 
sample 6. A characteristic of individuals 
from these samples is a light stripe with 
indistinct borders lying in the dorso- 
lateral dark field and extending from 
the ear to the groin (Fig. 23 C and D). 
This stripe is usually continuous but, if 
broken, the interspaces are much nar- 
rower than the remnants of the stripe. 
The stripe is white or blue-white in 
males, pale brown in females and is nar- 
rower and less distinct in females. The 
ventrolateral field contains vertically 
oriented bars or blotches on a dark 
background in adult males. Some males 
in sample 5 show a tendency for the 
blotches in the dorsolateral field to be 
continuous with those in the ventrolat- 
eral field. The ventrolateral field of 
females is mottled. Females in samples 
6 and 28 occasionally have a secondary 
dark stripe between the dorsolateral 
light stripes and the lighter brown area 
of the middorsal field, the same as de- 
scribed for individuals from samples 7 
and 8. 

Adult male A. wulnhta in samples 
1 and 2 are similar in pattern to those 
just described, but the dorsolateral dark 
field contains a series of large, quad- 
rangular, white or blue-white blotches 
which are two or three times as wide as 
the interspaces. These blotches are 
never fused into a solid stripe. The 
dorsolateral dark field of females con- 
tains similar, but less prominent 
blotches. The dorsolateral fight stripe, 
which is retained in its entirety in some 
adult females, terminates at one-half to 
two-thirds of the distance from occiput 
to groin in juvenile males and may be 
present only on the neck in adults. 

Adult males from samples 22, 23 and 
48 are similar to those of samples 1 and 
2, except that the blotches in the dorso- 
lateral dark field may be subequal in 
width to the interspaces. Samples 24 
and 25 contain adult males with dorso- 
lateral blotching as in samples 22 and 
23, but also contain some males in 



which the blotches are fused into a con- 
tinuous stripe as described for samples 
4, 6 and 28. In addition, there is a 
greater tendency for the dorsolateral 
light stripe to be retained in males from 
samples 24 and 25 than in males from 
samples 22 and 23. 

With few exceptions, other samples 
of Ameiva undulata from Pacific drain- 
age localities in Mexico present a variety 
of lateral patterns ranging from pres- 
ence of blotches in the dorsolateral dark 
field and spots, blotches or bars in the 
ventrolateral field to having the entire 
flanks barred or a combination of both 
patterns. No one pattern type prevails 
among males of any one sample. Adult 
males in sample 44 (El Zamorano and 
vicinity, Depto. Francisco-Morazan, 
Honduras) are unusual in that both 
males and females possess numerous 
(12-13 per side) small blotches in the 
dorsolateral dark field. These blotches 
are tear-drop shaped, and some extend 
ventrally (Fig. 23 G and H). This pat- 
tern is not apparent elsewhere. 

Samples 9-14, 16-19, 21, 34, 39-43 
and 46 are characterized by a lateral 
pattern of white or blue-white bars 
which extend from the dorsolateral light 
stripe or just beneath it to or almost to 
the enlarged ventral scutes ( Figs. 21 C 
and 23 A; Taylor, 1956, Fig. 67). The 
bars show little or no differentiation 
into dorsolateral and ventrolateral com- 
ponents. The dorsolateral dark field is 
usually present in adult males, but it is 
difficult to define because it grades grad- 
uallv into the ventrolateral field. In 
such males the dorsolateral and ventro- 
lateral light stripes are retained only 
anteriorly and the ventrolateral light 
stripe may be completely lacking in 
large adults. Sexual dimorphism is 
marked. With the exception of those 
from Isla del Maiz Grande (sample 21), 
adult females retain a juvenile pattern 
with the addition of blotches in the 
dorsolateral dark field (Fig. 21 D-F). 
Adult females from Isla del Maiz 
Grande resemble adult males in pattern 



56 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



TABLE 2. — Total number of lateral bars 
for some samples of male Ameiva undulata. 
Arranged in descending order according to 
sample mean. 



Sample 
Number 


X 


S.D. 


N 


Range 


46 


29.7 


1.11 


7 


28-31 


12 


25.4 


3.47 


9 


20-31 


9 


24.6 


3.58 


5 


20-28 


10 


24.3 


3.20 


14 


19-29 


34 


21.6 


1.67 


5 


20-24 


42 


21.0 





4 


21 


21 


20.0 


1.95 


20 


16-23 


41 


20.0 


1.81 


12 


16-22 


43 


19.8 


1.34 


7 


18-22 


18 


19.6 


2.25 


16 


16-23 


14 


19.5 


1.77 


8 


17-22 


13 


19.4 


1.81 


9 


16-21 


39 


18.6 


1.78 


10 


17-22 


40 


18.3 


1.97 


6 


16-21 


17 


17.8 


1.28 


8 


16-20 


19 


16.8 


1.92 


5 


15-20 



(Fig. 23 A-B), but not in color. The 
total number (sum of number on both 
sides) of vertical bars of males varies 
considerably among these samples 
(Table 2). The largest sample means 
are those of samples on the Yucatan 
Peninsula and Isla Mujeres. The sample 
from Isla Mujeres (sample 46) has a 
mean of 29.7, the largest recorded. 
Lowest sample means occur in the two 
northern coastal samples from Hon- 
duras (samples 17 and 19), with means 
of 17.8 and 16.8, respectively. Males 
from samples 9-14, 17-19, 34 and 46 may 
be differentiated from barred males 
from other samples in that they have 
little or no secondary spotting or 
blotching between the bars in the ven- 
trolateral field. Taken as a whole, 
barred males from other samples have 
such markings to a greater degree, al- 
though individuals from these samples 
may show few secondary markings. 

The general ontogenetic changes ex- 
hibited by the barred populations of 
Ameiva undulata are illustrated in Fig- 
ure 21. Neill (1961, 1965), reported that 
young undulata from parts of British 



Honduras lack lateral light stripes, the 
ontogenetic progression being from un- 
striped hatchlings through light-striped 
subadults to barred adults. Presumably 
this refers to ontogeny of males only. 
Neill ( 1965 ) also called attention to 
two small specimens (MCZ 71623-24) 
from Augustine, Cayo Dist., British 
Honduras. MCZ 71623 has four light 
lines on each side, whereas MCZ 71624 
has three. Augustine is in the ecolog- 
ically distinctive Mountain Pine region 
of British Honduras. 

Among samples of Ameiva undulata 
there is considerable geographic varia- 
tion in amount of dark pigment scat- 
tered as flecks or blotches in the mid- 
dorsal area. These dark markings usual- 
ly take the form or more-or-less paired 
or alternating, paravertebral spots, al- 
though in heavily pigmented specimens 
a reticulate pattern may be evident. 
The latter is especially evident in speci- 
mens from Isla del Maiz Grande (Fig. 
23 A-B). Both sexes may exhibit such 
markings. Although they are sometimes 
evenly distributed, the markings are of- 
ten restricted to or heaviest on the pos- 
terior one-half or two-thirds of the 
back. I have not attempted to quantify 
this aspect of pattern but feel justified 
in making the generalization that the 
samples along the Pacific slopes from 
southeastern Chiapas, Mexico through 
El Salvador (samples 33, 35-38) and 
northward into Honduras south of the 
northern coastal ranges (samples 16, 20 
and 44) are the most heavily marked of 
the mainland populations sampled. 
Specimens from Isla del Maiz Grande 
(sample 21) have very heavy middorsal 
marking, as noted above, but this 
marking takes a different form, appear- 
ing to be a continuation of the lateral 
pattern. Samples 39 and 40 are variable 
with respect to this character and sam- 
ples 41-43 show very light markings. A 
few specimens of Ameiva undulata are 
available from the Meseta Centrale of 
Costa Rica. This population was named 
by Cope (1894) as Cnemidophorus 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



57 




Fig. 24. Aiijeiva undulata (Holotype of 
Cnemidopliorus amivoides Cope; AMNH 
16316) from La Caipintera, Cartago Prov.?, 
Costa Rica, illustrating the heavy blotching in 
the middorsal area and the divided inter- 
parietal; a male, SVL 64 mm. 

amivoides (see Remarks). These speci- 
mens all show a high degree of marbling 
or blotching in the middorsal region 
(Fig. 24), a pattern sharply divergent 
from that of other populations of undu- 
lata at the southern end of its range. 
Further study of this population is de- 
sirable. Samples other than those spe- 
cifically mentioned above show variable 
markings from light to moderate in 
extent. 

There is little geographic variation 
with respect to color other than that di- 
rectly associated with pattern. Studies 
of such variation are hampered by a 
paucity of good color-in-life descrip- 
tions. Such descriptions are available 
for samples from Michoacan, Mexico 
(Duellman, 1961), the Yucatan Penin- 
sula (Maslin, 1963; Duellman, 1965a), 
British Honduras (Neill, 1965) and 
Chiapas (Alvarez del Toro, 1960). 
Other descriptions have been gleaned 
from my own field notes and those of 
William E. Duellman. 

The most striking aspect of geo- 
graphic variation in color has to do with 
color in the gular region of adult males. 
As noted for Ameiva festiva and quad- 
rilineata, the gular coloration of males 
may be associated with reproductive 
condition. In many specimens through- 
out the range, gular coloration (and 
ventral coloration in general) is white 
or cream. In other specimens, the en- 



tire venter is pale blue. A number of 
individuals, however, show either red 
or yellow pigment in the gular region 
and over the anterior part of the chest 
and ventral surfaces of the forelimbs. 
The red coloration ranges from orange 
or brick-red to scarlet. The yellow pig- 
ment is a bright lemon yellow. Whether 
one color precedes the other in a dy- 
namic system of change is not known, 
but I have observed no specimens in- 
termediate in coloration. The specimens 
upon which this discussion is based 
were collected during June, July or 
August between 1959 and 1966 by a 
number of collectors. Yellow pigment 
has been observed to the exclusion of 
red at only three localities in Mexico: 
along the Rio Frio, Tamaulipas; Finca 
Orizaba, Chiapas; and Tapanatepec, 
Oaxaca. At some localities, individuals 
may have either red or yellow pigment 
in the gular region. Thus, Duellman 
(1965a) reported that about half of his 
sample from Piste, Yucatan had orange 
throats, whereas half had yellow throats, 
corroborating an observation by Maslin 
(1963). Other localities from which 
both colors have been reported are 
Felipe Carillo Puerto, Quintana Roo 
(Duellman, 1965a), British Honduras 
( Neill, 1965 ) and the Tepalcatepec Val- 
ley of Michoacan (Duellman, 1961). 
Specimens from a number of other 
localities in Veracruz, Tabasco, Cam- 
peche, Yucatan, Quintana Roo, Chiapas, 
Jalisco, Honduras and Isla del Maiz 
Grande, Nicaragua had only red- 
throated individuals. 

There seems to be some geographic 
variation in coloration of the middorsal 
area. This area is iridescent moss- 
green anteriorly grading to yellowish- 
brown posteriorly in specimens col- 
lected by me at La Ceiba, Coyoles and 
Trujillo, Honduras (samples 17-19, re- 
spectively). It is reported as "olive" 
for specimens from British Honduras 
(Neill, 1965) and Chiapas (Alvarez del 
Toro, 1960). Specimens from the Te- 
palcatepec Valley, Michoacan (Duell- 



58 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



man, 1961); Piste, Yucatan (Maslin, 
1963); along the Rio Frio, Tamaulipas; 
16 km. SW Frontera, Tabasco; Mapas- 
tepec, Chiapas; vicinity of Melaque, 
JaHsco; El Zamorano, Depto. Francisco- 
Morazan, Honduras (personal observa- 
tion) have brown middorsal pigmenta- 
tion. Few descriptions of lateral ground 
color of living lizards are available, but 
there may be considerable geographic 
variation for this character as well. 

Ecological Notes. — Despite its broad 
range and relative abundance in many 
parts of that range, there have been no 
thorough ecological investigations of 
Ameiva undulata. Literature on this 
subject is limited to miscellaneous ob- 
servations. Thus, Stuart (1951) found 
undulata altitudinally distributed to 
about 2000 m in the "moderately hu- 
mid" region near Panajachel, Depto. 
Solola, Guatemala, but commented that 
in more moist areas the altitudinal limit 
is reached about 400 m lower. Else- 
where, the species seems to occur below 
1500 m, and its vertical distribution may 
be limited to below the level of occa- 
sional frosts (1200-1500 m, depending 
upon humidity), a factor considered by 
Stuart (1966) to be of possible bio- 
logical significance. 

Throughout most of its range, 
Ameiva undulata is found in forest or 
forest-edge situations (Alvarez del 
Toro, 1966; Duellman, 1965a; Echter- 
nacht, 1968; Rand, 1957; Stuart, 1935, 
1950, 1954b, 1958) or, in drier regions, 
dense thickets and scrub (Maslin, 1963; 
Schmidt and Stuart, 1941; Stuart, 1948). 

Stuart (1950), Duellman (1966), 
and Echternacht (1968) commented on 
the sciophilic nature of Ameiva iindu- 
lata, although I pointed out that even 
the more shade-tolerant species of 
Am^eiva may utilize open areas in the 
absence of competition from more he- 
liophilic macroteiids like Cnemidopho- 
rus. Stuart (1950) reported undulata 
to be abundant in the grassland and ri- 
parian habitats of Depto. Alta Verapaz, 
Guatemala, and Echternacht ( 1968 ) 



found the species to be common in open 
situations near Panajachel, Depto. So- 
lola, Guatemala. Although the lizards 
shun the open beach, I found the 
species occupying clearings, paths and 
other open areas as well as thickets and 
piles of debris in the coconut groves of 
Isla del Maiz Grande, Depto. Zelaya, 
Nicaragua. In northern Honduras, 
western Nicaragua, northwestern Costa 
Rica, and in those parts of Mexico with 
which I am familiar, undulata occurs in 
shaded areas; the more open parts of 
the habitat are occupied by various 
species of Cnemidophorus. 

Virtually nothing has been reported 
on the reproduction of this species of 
Ameiva. As I have remarked elsewhere, 
research on this topic may reveal the 
sequence and basis for changes in gular 
pigmentation of adult males. 

Stuart (1958) collected 56 juveniles 
and only four adults of Ameiva undu- 
lata at Tikal, El Peten, Guatemala, be- 
tween February and May, 1956, but he 
cited a collection made by Gosner at 
the same locality at unspecified dates in 
1947 and 1949 that contained a "fine 
series of adults." This, along with my 
own observations, suggests that adult 
and subadult Ameiva may have differ- 
ent seasonal activity periods. However, 
there have been no observations of a 
quantitative nature to substantiate this. 

Remarks. — I have examined four 
specimens of Ameiva undulata (ANSP 
15438-15441) from "Wounta" (= Hua- 
unta) Haulover, Depto. Zelaya, Nica- 
ragua. This locality is at least 290 kilo- 
meters from other known localities for 
the species. Duellman (1958) exam- 
ined three specimens (ANSP 15445- 
15447) of the snake Leptodeira annu- 
lata rhomhifera Giinther from the same 
locality, far removed from the re- 
mainder of the range of the species. Al- 
though it is possible that a snake might 
have been carried to Huaunta by man, 
it is unlikely that a moderately large, 
active lizard could be so transported. 
Ameiva undulata shuns deep forest, and 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



59 



it is unlikely to have arrived at Huaunta 
along rivers from the west, but it is pos- 
sible that the species exists in isolated 
or semi-isolated populations along the 
Atlantic coast of Nicaragua northward. 
An alternative hypothesis would be that 
the A7newa and Leptodeira populations 
at Huaunta are relict and that a more 
careful survey of the region would un- 
cover other such species as well. This 
part of Central America certainly de- 
serves more attention from biologists 
than it has received. 

Such an explanation does not apply 
for records of Ameiva undidata from 
Panama (Brennan and Yunker, 1966). 
All specimens so reported are lepto- 
phrys. 

Taylor (1956) placed Cnemidopho- 
rus amivoides Cope from the Meseta 
Central of Costa Rica in the synonymy 
of Ameiva undidata, and I see no rea- 
son to dispute this judgment. I have 
called attention to the unusual color 
pattern of specimens from the Meseta 
Central (Fig. 24), but so few are avail- 
able that further comment is unjustified 
at this time. 

Dunn (1940a) and Villa (1968) in- 
cluded the population of Ameiva on the 
Islas del Maiz, Depto. Zelaya, Nica- 
ragua (sample 21) in the species undu- 
lata. The population was described as 
a subspecies of festiva by Barbour and 
Loveridge (1929a). I have substan- 
tiated the assignment of this population 
to undulata ( Echternacht, 1970). 

Ameiva chaitzami Stuart 

Ameiva chaitzami Stuart, Proc. Biol. Soc. 
Washington, 55:143-145, 1942 [Holotype: 
UMMZ 90638. Type Locality: "Along 
Cahabon-Languin trail about 2 km north 
of Finca Canihor . . . , Alta Verapaz, 
Guatemala." Collector: L. C. Stuart]. 
Stuart, Misc. Publ. Mus. Zool., Univ. Mich- 
igan No. 122, p. 77, 1963. Echternacht, 
Breviora, 354: 7, 1970. 

Ameiva undidata stuarti — Stuart, Occas. Papers 
Mus. Zool, Univ. Michigan No. 471, p. 21, 
1943. 

Ameiva undulata thomasi Smith and Laufe, 
Univ. Kansas Sci. Bull. 31(2):47-50, Pi. 
lA, 1946 [Holotype: FMNH 100006. Type 



92» 



' "\ — 



5° 



I 



',• 




s 



10 50 

H 1 

KILOMETERS 



92° 



90° 



89° 



88° 



Fig. 25. Map showing locality records of 
Ameiva chaitzami. Samples used in statistical 
analyses were ( 1 ) Comitan, Chiapas, Mexico, 
(2) Vicinity of San Antonio Huista, Depto. 
Huehuetenango, Guatemala, (3) combined 
sample of those specimens originally desig- 
nated by Stuart (1942) as holotype and para- 
types from the Languin-Cahabon road near 
Finca Canihor, Depto. Alta Verapaz, Guate- 
mala, and specimens from the vicinity of 
Poptun, Depto. El Peten, Guatemala, later 
collected by Stuart and identified as A. 
chaitzami. 



Locality: La Libertad, Chiapas, Mexico, 
near Rio Cuilco where it crosses Guate- 
malan border. Collector: H. D. Thomas]. 
Smith and Taylor, U. S. Natl. Mus. Bull. 
199, p. 173, 1950. 

Distribution. — Known from the val- 
leys of the upper tributaries of the Rio 
Grijalva in Chiapas, Mexico and west- 
central Guatemala, from along the Lan- 
guin-Cahabon trail near Finca Canihor, 
Alta Verapaz, Guatemala, and near 
Poptun, El Peten, Guatemala (Fig. 25). 

Diaanosis. — Ameiva chaitzami can 
be distinguished from other Middle 
American congeners by the following 
combination of characters: Small size 
(maximum observed SVL of males 85 
mm, of females 75 mm); central gular 
scales enlarged, in longitudinal arrange- 
ment; gradual reduction in size of scales 
radiating outward from central gular 
scales; prefrontal scales in contact with 



60 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



postnasal scales; three parietal scales 
(four if median parietal scale divided); 
a transverse row of abruptly enlarged 
mesoptychial scales; usually eight trans- 
verse rows of ventral scutes; relatively 
narrow middorsal stripe (mean PV =: 
37.1); no narrow, well-defined vertebral 
stripe; usually a dark secondary stripe 
medial to the dorsolateral light stripes; 
dorsolateral blotches of males fused to 
the dorsolateral light stripe such that 
the latter has a well-defined dorsal bor- 
der and an irregular ventral border. 

Adult male Ameiva chaitzami can 
be distinguished from other Ameiva 
readily on the basis of pattern. Juvenile 
individuals and females are difficult to 
differentiate from some undidata, espe- 
cially those from the coastal lowlands 
of eastern Veracruz, Tabasco, Chiapas 
and extreme western Campeche, Mex- 
ico. However, there are no signs of 
intergradation or hybridization of chait- 
zami with other species of Ameiva. The 
only Ameiva with which chaitzami 
comes in contact is the large form of 
undulata having a color pattern as 
shown in figure 21, and there is no dif- 
ficulty in distinguishing the two species. 

Description. — Maximum observed 
SVL 85 mm for males, 75 mm for fe- 
males; supralabials 13-16 (14.2); infra- 
labials 9-13 (10.1); supraoculars 6-8 
(6.2); COF value 2; COP value 10-13 
(11.6); SO-SC value 2-8 (2.3); loreals 
2-4 (3.7); nostril in prenasal-postnasal 
suture; prefrontals in contact with post- 
nasals; central gular scales enlarged, 
longitudinal in arrangement; GAB 83- 
134 (108.6); PV 26-51 (37.1); PV/GAB 
0.27-0.42 (0.34); GOR 174-243 (203.6) 
for males, 171-255 (205.1) for females; 
GAB/GOR 0.44-0.65 (0.54); SAT 20-24 
(21.2); total preanal scales 5-9 (6.8); 
terminal preanal scales 1-3 (2.0); one 
row of enlarged prebrachials; one or 
two rows of enlarged postbrachials; en- 
larged preantebrachials in two rows or 
two rows proximally and one row dis- 
tally; prefemoral scales enlarged, four 
(usually) or three rows proximally re- 




FiG. 26. Head and preanal scutellation of 
Ameiva chaitzami (UMMZ 90643, paratype, 
from along the Cahabon-Langiiin trail about 
2 km N Finca Canihor ) : ( A ) Lateral view of 
head, ( B ) Dorsal view of head, ( C ) Ventral 
view of head, X 2. (D) Preanal region, 
X 2.7. 

ducing to three (usually) or two rows 
distally; infratibials enlarged, in two 
rows; femoral pores 30-41 (35.0) for 
males, 26-36 (32.7) for females; sub- 
digital lamellae 24-33 (28.6); longitudi- 
nal rows of ventral scutes 27-32 (30.0); 
transverse rows of ventral scutes 8-10 
(8.1). For details of scutellation see 
figure 26. 

Color and Pattern in Alcohol. — De- 
scriptions of specimens in preservative 
have been presented by Stuart ( 1942 ) 
and Smith and Laufe ( 1946 ) . Juvenile 
males possess a broad middorsal field 
bounded laterally by narrow, white dor- 
solateral stripes which originate just be- 
hind the eye and extend well onto the 
tail. Between the blue-gray or gray- 
brown middorsal pigment and the dor- 
solateral light stripes there are often 
secondary stripes, velvet black in color, 
which extend from the occiput to the 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



61 



base of the tail. The medial border of 
the secondary stripe is irregular. The 
niiddorsal field may be flecked witli 
black or be nearly uniform in colora- 
tion. A white ventrolateral light stripe, 
which may be complete or interrupted, 
extends from the posterodorsal corner 
of the ear to the groin, beginning again 
lieliind the leg and terminating on the 
tail. A white postfemoral stripe is con- 
tinuous with the ventrolateral light 
stripe behind the leg. The black dorso- 
lateral dark field is prominent and con- 
tains faint, light-colored blotches or 
bars. The ventrolateral field contains 
wliitc mottling or bars on a black back- 
ground. The same pattern is evident on 
dorsal surfaces of the limbs, but the 
mottling is more pronounced. Ventral 
surfaces are cream or white in color. 

Juvenile females resemble juvenile 
males except that there are no light- 
colored blotches or bars in the dorso- 
lateral dark field and the ventrolateral 
and limb mottling is less well de- 
veloped. 

Through ontogeny (Fig. 27) the 
light ventrolateral and dorsal limb mot- 
tling or barring becomes much more 
pronounced and the ventrolateral light 
stripe becomes interrupted such that it 
merges with the ventrolateral mottling. 
The dorsolateral blotches become white 
or blue-white in color and fuse with the 
dorsolateral light stripe in males, giving 
that stripe an irregular ventral border. 
Adult females resemble juveniles of 
both sexes in color and pattern, ap- 
parent ontogenetic changes being slight. 
Color and Pattern in Life. — There 
are no descriptions of the coloration 
of living Ameiva chaitzami, although 
Stuart's (1942) description of the holo- 
type may have been written before ex- 
tensive fading had occurred in preserva- 
tive. He stated that the middorsal field 
and top of the head are olive-brown and 
that the legs and arms are olive-brown 
above with black and bluish-white 
mottling. The anterior surfaces of the 
thighs are said to be black with blue 



spots, the venter l:)hiisli and the head 
blue mottled with black. There is no 
mention of yellow or red pigment in tlie 
gular region of this species, the only 
Middle American Ameiva with which I 
have had no field experience. 

Geographic Variation. — The follow- 
ing characters show no significant dif- 
ferences among sample means: Femoral 
pores of males, number of supralabials, 
number of infralabials, number of ter- 
minal preanals, number of transverse 
rows of ventral scutes, SAT and COF, 
COP and SO-SC values. 

The three samples of Ameiva chait- 
zami (Fig. 25 ) were collected in eastern 
Chiapas, Mexico (Sample 1), western 
Guatemala (2) and central to eastern 
Guatemala (3). Samples 1 and 2 are 
much closer geographically than either 
is to sample 3 and sample 1 lies slightly 
northwest of sample 2. This geographic 
arrangement is reflected in the results 
of STP analysis of characters which 
show significant differences among 
means. Five characters show samples 1 
and 2 as constituting one subset, sample 
3 a second subset significantly different 
from the first. These five characters are 
numbers of loreals and subdigital lamel- 
lae, GAB /COR, numbers of longitudi- 
nal rows of ventrals and total preanal 
scales. For two characters, GAB and 
PV/GAB, samples 2 and 3 form a sub- 
set distinct from that containing only 
sample 1. For both PV and GOR of 
females, all three sample means fall into 
separate, significantly distinct subsets. 
This is true of samples 1 and 2 for num- 
ber of femoral pores of females and 
GOR of males; sample 3 is too small for 
inclusion in STP analysis. Only for one 
character, number of supraoculars, is 
there overlap of sample means: there is 
a gradient from sample 3 through sam- 
ple 2 to sample 1, samples 3 and 2 con- 
stituting one subset and samples 2 and 
1 another. 

Because of the inadequate number 
of samples and the geographic localities 
from which the samples were drawn, 



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ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



63 



detection of clinal trends must await 
accumulation of additional large sam- 
ples as the range of the species becomes 
more fully known. 

There seems to be no notable geo- 
graphic variation in color, pattern or 
arrangement of gular scales among the 
three available samples. A comparison 
of color patterns of equal-sized (68 
mm) males from samples 1 and 3 may 
be seen in Figure 27 (B and G, re- 
spectively ) . 

Ecological Notes.— Stuart ( 1942; 
personal communication) provided the 
only insight into the ecology of Ameiva 
chaitzami. He is of the opinion that the 
species is restricted to pine savannas on 
the lowlands of central Guatemala 
where its population density may be 
kept at a low level by regular burning- 
off of ground cover. The species is 
found with Ameivo tmdulata, but never 
with Cnemidophorus. Other Middle 
American species of Ameiva are sym- 
patric with species of Cnemidophorus 
over at least part of their range. Stuart 
(1942) mentioned a chaitzami with a 



SVL of 66 mm having "well formed 
eggs." 

Smith and Laufe (1946) stated that 
these lizards ". . . probably occur in all 
the dry, hot valleys of the upper tribu- 
taries of the Rio Grijalva in the interior 
of Chiapas and of western central 
Guatemala." 

Remarks. — Less is known about the 
biology of Ameiva chaitzami than of 
any other Middle American Ameiva. 
The overall range of the species has yet 
to be defined. 

The divided interparietal (median 
parietal), which was used by Stuart 
( 1942 ) to diagnose Ameiva chaitzami, 
is a variable character found through- 
out the genus in Middle America with 
varying frequency ( Echternacht, 1970). 
Likewise, I have found the lateral en- 
largement of the gular scales cited by 
Smith and Laufe ( 1946 ) to be an unre- 
hable character. Color pattern and size 
remain the most useful characters for 
distinguishing this probable sibling spe- 
cies from its closest relative, undulata. 



DISCUSSION 



INTRASPECIFIC RELATIONSHIPS 

In my opinion the Ameiva of Mid- 
dle America belong to three natural 
groups the relationships of which will 
remain obscure until their affinities with 
South American representatives of the 
genus are known. The three groups 
contain one, two and three species ( Fig. 
28). 



Fig. 27. Ontogenetic change in pattern of 
Ameiva chaitzami. A-D and F, UMMZ 94905, 
E, UMMZ 94901, all from the vicinity of 
Comitan, Chiapas, Mexico. Field Tag Num- 
bers and SVL are: Males, (A) 473, 45 mm, 
(B) 482,68 mm, (C) 461, 73 mm. Females, 
(D) 483, 55 mm, (E) 365, 68 mm, (F) 501, 
75 mm. (G) For comparison, UMMZ 90640, 
a male paratype 68 mm SVL from along the 
Cahabon-Languin trail about 2 km N Finca 
Canihor, Alta Verapaz, Guatemala. 



Ameiva ameiva, the only species in 
Group I, can be distinguished from the 
others on the basis of a number of 
scale and pattern characteristics. Chief 
among these are the presence of five 
parietal scales, 10-12 enlarged ventral 
scutes, a large number of scales around 
the tail, high GAB and GOR counts and 
a pattern which emphasizes spots in- 
stead of stripes in adults. Additionally, 
ameiva is the only species included in 
the present study having the bulk of its 
range in South America. The Pana- 
manian portion of that range is but a 
minor and disjunct part of the whole. 
Group I is farther removed phylogenet- 
ically from Groups II and III than these 
are from each other. 

Group II includes Ameiva festiva 
and leptophrijs. Both species are essen- 
tially lower Central American in dis- 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 




Fig. 28. Relationships of the Middle American 
species of Ameiva. Group I: A, A. ameiva. 
Group II: F, A. f estiva; L, A. leptophrys. 
Group III: Q, A. quadrilineata; C, A. chait- 
zami; U, A. undulata. Lengths of the lines 
and positions of branches do not reflect a 
time scale. 

tribution. Ameiva festiva occurs in Co- 
lombia, and leptophrys is expected 
there. The group is characterized by 
the relatively high number of femoral 
pores and a tendency for the enlarged, 
posterior scales of the head to become 
disrupted (Fig. 7). Disruption of head 
scales is more noticeable in leptophrys 
than in festiva but is a more commonly 
encountered feature of these species 
than of other Middle American species 
of Ameiva. 

Group III consists of Ameiva quad- 
rilineata, chaitzami and undulata. The 
group is entirely Middle American in 
distribution, and is characterized by 
similarities in pattern and scale charac- 
ters. The pattern emphasizes stripes, 
rather than spots, and features a broad, 
middorsal field without a vertebral 
stripe. Prefrontal and postnasal scales 
are in contact. Except for an occasional 
longitudinal division of the interparietal 
scale, there is little disruption of pos- 
terior scales of the head. Ameiva chait- 
zami is very closely related to undulata, 
from which it is distinguishable pri- 
marily on the basis of the color pattern 
of males. However, the two species are 
sympatric in central Guatemala. Clari- 



fication of the relationships of chait- 
zami, its range and biology remain 
problems of major importance. 

Elucidation of the relationships of 
the Middle American species of Ameiva 
with those of South America may show 
that Groups II and III are much more 
closely related than I have indicated. 
These may constitute a single group. I 
should emphasize that these three 
groups of Middle American Ameiva are 
not the equivalents of the species 
groups of Cnemidophorus established 
by Burt ( 1931 ) . Nevertheless, it seems 
apparent that my Group I would belong 
to a difterent species group (sensu 
Burt) than the other species of Middle 
American Ameiva, regardless of whether 
these other species are found to belong 
to one or several species groups. The 
relationships indicated without substan- 
tiating comment by Barbour and Noble 
(1915) are suspect. The taxonomic im- 
portance of extensive division of head 
scales in Ameiva bridgesi Cope, orcesi 
Peters and septemlineata Dumeril and 
Dumeril (Peters, 1964) and the rela- 
tionships of these species to leptophrys 
and festiva require further study. The 
relationships of niceforoi also need clari- 
fication ( Echternacht, 1970). 

INTERSPECIFIC PATTERNS IN 
GEOGRAPHIC VARIATION 

Assessing interspecific trends in vari- 
ation is considerably more difficult than 
determining intraspecific trends. This is 
so for two reasons: 1) The narrow lin- 
earity of lowland Middle America vir- 
tually dictates that clines will run paral- 
lel to the coastlines along a northwest 
to southeast axis and 2) samples of two 
or more species from the same locality 
and of sufficient size for statistical anal- 
ysis are rarely available. Samples from 
only ten localities where sympatric oc- 
currence of two or more species was 
found were large enough for analysis 
(Table 3), and only three of the six 
Middle American species of Ameiva 
were present in more than one sample. 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



65 



TABLE 3. — Localities from which samples of two or more species were available for statistical 

analysis. 



Locality 



Species 



Costa Rica: Limon: Limon, Los Diamantes and vicinity 

Costa Rica: Limon: Tortuguero 

Costa Rica: Puntarenas: Golfito Region 

Costa Rica: Puntarenas: Osa Peninsula 

Guatemala: El Peten: Piedras Negras 

Honduras: Northern Coastal Localities 

Panama: Bocas del Toro: Almirante and vicinity 

Panama: Canal Zone 

Panama: Isla Escudo de Veraguas 

Panama: Eastern Darien 



festiva. 


quadrilineata 


festiva. 


quadrilineata 


festiva. 


quadrilineata, leptophrys 


festiva. 


(juadiilineata, leptophrys 


festiva. 


tmdidata 


festiva. 


itudulata 


festiva. 


quadrilineata 


ameiva 


festiva, leptophrys 


festiva. 


quadrilineata 


festiva. 


leptophrys 



Ameiva undtdata was present with fes- 
tiva in but two samples, and festiva was 
found with quadrdineata in two. Al- 
though no rigorous statistical analysis 
was attempted, some parallel trends are 
evident. For seven characters (infra- 
labials, supraoculars, COP, GAB, GOR, 
subdigital lamellae and scales around 
the tail) an increase or decrease in 
sample mean for festiva was paralleled 
by a like shift in mean for the same 
character for tindulata when the two 
localities of sympatry (Piedras Negras, 
El Peten, Guatemala, and northern Hon- 
duras) were compared. Similar trends 
were noted for festiva and quadrdineata 
for ten characters (GAB, GOR, GAB/ 
GOR, longitudinal rows of ventral 
scutes, transverse rows of ventral scutes, 
femoral pores, subdigital lamellae, total 
preanal scales, temiinal preanal scales, 
and scales around the tail) when step- 
wise pair comparisons of samples were 
made between localities beginning with 
Tortuguero in Costa Rica and running 
through the Canal Zone to Darien, 
Panama, and thence to the Golfito and 
Osa Peninsula regions of Costa Rica. 
Perfect agreement was noted only for 
GOR, but all other characters men- 
tioned showed parallel mean-shifts for 
at least two-thirds of the possible pair 
comparisons. Similar parallel changes 
were noted for festiva and leptophrys 
( supralabials, infralabials, COF, COP, 
transverse rows of ventrals and scales 



around the tail with COP and COF 
showing perfect agreement) and lepto- 
phrys and quadrilineata (infralabials 
and supraoculars). Such parallel shifts 
in sample means would tend to indicate 
that these characters are responding in 
similar ways to environmental differ- 
ences between sample localities. Con- 
siderable additional study will be neces- 
sary before any light will be shed on the 
question of which environmental factors 
are most important in determining the 
direction of shift of sample mean for a 
character from one sample locality to 
another. Because only increase or de- 
crease in the value of the mean were 
noted from sample to sample, and not 
the magnitude of the increase or de- 
crease, it is probable that some shifts 
are only the result of small sample size 
and that additional data would show 
that there has been, in fact, no shift 
at all. 

An important, although likewise un- 
explained, trend is that of insular popu- 
lations of Ameiva to have higher mean 
GAB and/ or GOR counts than main- 
land populations (Table 4). Samples 
are available from Isla Mujeres, Quin- 
tana Roo, Mexico (undidata), Isla del 
Maiz Grande, Zelaya, Nicaragua (undu- 
lata), Isla Escudo de Veraguas, Panama 
(festiva, quadrilineata) and Isla San 
Jose, Archipelago de las Perlas, Panama 
(ameiva). Samples from the Osa Penin- 
sula, Puntarenas, Costa Rica of festiva, 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



TABLE 4. — Relati\'e positions of island samples with respect to mean Granules Around the 
Body (GAB) and Granules Occiput to Rump (GOR) when sample means are ordered from largest 

to smallest. 





Total Number 






Relative 


Position 


Species 


of Samples 




GAE 


; 


GOR 


ameiva 


7 




1 




2*' " 1" 


festiva: Isla Escudo de Veraguas 


20 


1" 




1" 


V 1" 


fcstica: Osa Peninsula 


20 


4" 




4" 


15" 16" 


k'pto))hnjs 


4 




2 




2 


quadrilineata: Isla Escudo de Veraguas 


10 




7 




5 


quadrilineata: Osa Peninsula 


10 




1 




1 


imdulata: Isla Mujeres 


45 




2 




10 


uudulata: Isla del Maiz Grande 


45 




1 




2 



" Males where sexual dimorphism is evident. 

" Females where sexual dimorphism is evident. 

'' Tied with sample 3 (See Appendix B) for this position. 



leptophrijs, and quadrilineata also have 
relatively high dorsal granule counts; 
the peninsula may be effectively an is- 
land as far as Ameiva are concerned 
inasmuch as it is separated from the 
mainland by a swampy area. It will be 
obvious from an examination of table 4 
that not all insular samples have high 
doi-sal granule counts, but the tendency 
is evident in a remarkable number of 
instances. There is no clear relationship 
between high GAB and GOR counts on 
the one hand and the ecology of the 
islands upon which the lizards are living 
on the other. It would seem that Amei- 
va adapted to life on dry islands would 
have fewer, but larger, dorsal granules 
than Ameiva of the same size living on 
wet islands as a selective strategy for 
prevention of excess water loss. Fewer 
and larger granules would mean less ex- 
posed skin between the granules 
through which water might be lost than 
more and fewer scales on animals of 
equal size. Such an hypothesis is con- 
traindicated by the observation that the 
islands from which samples of Ameiva 
are available range from very dry (Isla 
Mujeres) to very wet (Isla Escudo de 
Veraguas and the Osa Peninsula). It 
would be instructive to have samples of 
one species of Ameiva from a number 
of localities for which weather records 
are available and ranging from rela- 



tively dry to relatively wet. Such sam- 
ples are not presently available and, 
until they are, this problem is likely to 
remain unsolved. 

Sample mean snout-vent length 
(SVL) data are not presented in this 
report because collections are usually 
strongly biased in favor of large, bright- 
ly colored males. The relatively drab 
juvenile and adult female Ameiva are 
often overlooked by collectors who ob- 
tain Ameiva incidental to those genera 
they themselves are working on. A sea- 
sonal bias also acts to reduce the num- 
bers of hatchling Ameiva in collections 
in that, in areas having pronounced wet 
and dry seasons, reproduction may be 
temporally restricted with hatchlings 
present only at certain times of the year. 
Collections made at other times do not 
include hatchlings. All of the insular 
samples of Ameiva available to me were 
obtained in June, July, or August, and 
all are the result of but one or two field 
trips. With the exception of my own 
collection from Isla del Maiz Grande, 
all of these collections were made by 
individuals primarily interested in other 
genera. For these reasons, I have little 
faith in the mean SVL of samples used 
in this study as an estimate of the true 
population means. Nevertheless, it is 
apparent that the insular samples have 
generally higher mean SVL than main- 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



67 



TABLE 5. — Correlation between snout-vent length (SVL) and granules around the body 
(GAB) or granules from occiput to nunp (GOR) for five species of Middle American Ameiva. Cor- 
relation coefficient followed by sample size in parentheses. 



ametva 



festiva 



leptophrys quadrilineata undulata 



SVL-GAB Males 
SVL-GAB Females 
SVL-GOR Males 
SVL-GOR Females 



0.1101(112) 0.0897(243) 0.1345 ( 101 ) -0.0289 ( 180) 0.1606(409) 

-0.0403(133) 0.1531 (233) -0.1040(98) -0.0950(114) 0.2445(561) 

0.0737(114) 0.1857(268) 0.2262(188) 0.1629(507) 

-0.0888(138) 0.0682(264) -0.2283(126) 0.1550(448) 



land samples, making it necessary to ask 
the question whether GAB and GOR 
counts are in any way correlated with 
SVL. Correlation analysis was carried 
out on a species-by-species basis (Table 
5). There is no indication that either of 
the tsvo dorsal granule counts are highly 
correlated with SVL and the high mean 
GAB and GOR values of island samples 
is not an artifact resulting from biased 
estimates of sample mean SVL. 

HISTORY OF AMEIVA IN 
MIDDLE AMERICA 

There is no fossil record of macro- 
teiids in Middle America. Therefore, 
any discussion of the history of Ameiva 
in that region must lean heavily on 
what is known of present distribution 
patterns and on geological and faunal 
summaries relating to Middle America 
and northern South America. Geological 
theories concerning Middle America 
have undergone an evolution of their 
own over the past 50 years; geological 
data were summarized by Lloyd ( 1963 ) , 
Vinson and Brineman (1963), and Mal- 
donado-Koerdell ( 1964 ) . Duellman 
(1966), Savage (1960, 1966) and Stuart 
(1935, 1951, 1954a, 1954b, 1957, 1964, 
1966) provided thoroughly documented 
discussions of the ecology, origins, and 
history of the herpetofauna of Middle 
America, whereas Haffer ( 1967, 1970 ) 
presented an important assessment of 
the geologic-climatic history of the criti- 
cal Gulf of Uraba-northwestern Colom- 
bia region. Although the following dis- 
cussion is based in large part on these 
works, it is a highly speculative and 



personal view and should be read with 
that in mind. 

Dunn ( 1931 ) considered the family 
Teiidae to be a part of his South Amer- 
ican element, having its origins in South 
America with some genera invading 
Middle America and one, Cnemido- 
phorus, reaching North America. The 
South American element itself was de- 
rived from a generalized tropical Amer- 
ican herpetofauna which ranged over 
much of North and South America dur- 
ing the early Tertiary ( Savage, 1960 ) . 
Subsequent authors have followed this 
arrangement (Duellman, 1965b; Savage, 
1960, 1966; Schmidt, 1943; Stuart, 1957, 
1964), although Savage (1966) sug- 
gested that the ancestral stock of Cnemi- 
dophorus was isolated north of the 
Isthmus of Panama and that of Ameiva 
south of the Isthmus when the Pana- 
manian portal was open throughout 
most of the Tertiary. Northward move- 
ments of the South American element 
would have been possible after the 
closure of the portal in the late Pliocene 
or early Pleistocene (Lloyd, 1963; Mal- 
donado-Koerdell, 1964). As outlined by 
Stuart (1957) and Savage (1966, Fig. 
23), Ameiva, with the exception of A. 
ameiva, probably followed humid, low- 
land routes from South America into 
Middle America. Movement through 
the Pacific lowlands progressed only as 
far as the Golfito-Palmar-San Isidro del 
General region of Costa Rica. An At- 
lantic route was followed northward in 
Mexico, dispersal to the Pacific lowlands 
occurring at several points north of 
what is now Lake Nicaragua. Some 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



southward movement along the Pacific 
lowlands of Nicaragua and Costa Rica 
at a later time is indicated by the pres- 
ent distribution of A. tinchilota ( Fig. 
17). These routes may have been 
broader than at present because of low- 
ered sea levels, especially in areas 
where the continental shelf is extensive 
(Bennett, 1968). 

Of all of the Ameiva found in Mid- 
dle America, A. ameiva has made the 
smallest inroads there. Heatwole (1966) 
placed its arrival in the Upper Pliocene 
at the earliest. As an alternative hy- 
pothesis, Bennett (1968) suggested 
that the species dispersed into Panama 
during the period of human occupation 
of Darien. The A. ameiva of Panama 
resemble those of western Venezuela; 
Bennett suggested than an open corri- 
dor between the two areas for the 
length of time suggested by Heatwole 
( over one million years ) v^^ould have re- 
sulted in formation of clines. Subse- 
quent closure of the corridor would 
have isolated opposite ends of such 
clines, resulting in morphologically di- 
vergent populations. Other than a study 
emphasizing color pattern ( Miiller, 
1929), no detailed study of the geo- 
graphic variation of South American A. 
ameiva has been reported. Donoso- 
Barros ( 1968 ) characterized the west- 
ern population of the species in Vene- 
zuela as having a light-colored vertebral 
stripe. Such a stripe is evident in those 
A. ameiva from eastern Panama, but 
lacking in those from the western part 
of that country (Fig. 4). With the 
examination of additional samples, rem- 
nants of a cline as postulated by Ben- 
nett may be found for this and other 
characters (but see Remarks, p. 19). If 
remnants of a cline are found, it would 
indicate that entry of the species into 
Middle America occurred at a time in- 
termediate to the extremes suggested by 
Heatwole and Bennett. Recent evi- 
dence of alternating dry interglacial and 
humid glacial periods during the Pleis- 
tocene so convincingly presented by 



Haffer (1967, 1970) would allow for 
several intervals during which suitable 
conditions prevailed for the northward 
movements of A. ameiva. Haffer (1970) 
suggested that the humid forests of gla- 
cial times were forced southward dur- 
ing interglacials and during post-Pleis- 
tocene dry periods and that central and 
eastern Panama, as well as the Gulf of 
Uraba-lower Rio Atrato region of Co- 
lombia, were not heavily forested. Thus, 
it would have been possible for A. 
ameiva to have moved into Panama 
prior to the period of human occupancy 
of the Darien and yet not long enough 
for establishment of clines. Indeed, if 
Haffer's scheme is followed, the corridor 
between central Panama and Colombia 
would have been interrupted and re- 
established several times, mediating 
against formation of clinal variation. In 
any case, it is certain that the Pana- 
manian A. ameiva originated from a 
population of South American Ameiva 
similar to the present population of the 
species in western Venezuela. Dispersal 
into Middle America was contained by 
the availability of non-forested habitats. 
The species probably reached the is- 
lands of the Archipelago de las Perlas 
at a time when a land connection existed 
between these islands and the mainland 
during a period of lowered sea level 
(Bennett, 1968). 

With the exception of Ameiva amei- 
va already discussed, none of the species 
of Ameiva presently inhabiting Middle 
America would have required a non- 
forested corridor for northward disper- 
sal. Although the species show a graded 
dependence upon forested habitat, all 
are found in or at the edge of forested 
areas. Ameiva f estiva and leptoplinjs 
may, upon completion of studies now in 
progress, be found to be closely allied to 
the A. septemlineata — hridgesi — orcesi 
assemblage of northwestern South 
America. Ameiva leptophrys, with its 
present range limited to eastern Pana- 
ma and the Pacific slopes of western 
Panama and southeastern Costa Rica 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



69 



( Fig. 5 ) may have evolved ( along with 
several other species of reptiles and am- 
phibians) on insular isolates during the 
Middle Tertiary (Savage, 1966). This 
seems possible in view of both the pres- 
ent distribution of the species and its 
broad sympatry with festiva. The ap- 
parent absence of populations of Jepto- 
phrys on islands at present is a minor 
set-back to a theory of insular origin in 
that suitable habitat may have disap- 
peared from the islands in the relatively 
recent past, or that further collecting 
will reveal the presence of the species. 
Ameiva festiva occurs in both Mid- 
dle and South America ( Fig. 9 ) , as does 
A. ameiva, but festiva is continuously 
distributed over its continental range. 
Populations of festiva on the west and 
east ( or north ) slopes of the Andes have 
diverged markedly in color pattern ( Fig. 
12 ) . On this basis, the population found 
in the Choco of Colombia seems to have 
diverged relatively early from the an- 
cestral stock. It may possibly have been 
isolated in the Choco region during a 
glacial period during the Pleistocene. 
Haffer ( 1967 ) documented similar iso- 
lation in birds and fomially proposed 
the name "Choco-Refuge" for the area. 
I have seen no specimens intermediate 
in pattern between those of the Choco 
(Pattern Type C; see page 31 for de- 
scriptions of pattern types) and speci- 
mens from populations to the north and 
east. Because of similarity in color pat- 
tern of festiva from eastern Colombia 
and those of the San Bias and Golfito- 
Puerto Armuelles regions of lower Mid- 
dle America, it seems likely that the 
latter are derived from the former, 
rather than from the Chocoan popula- 
tion. The species moved northward, 
through lowland forests along the At- 
lantic slopes. The evolution of Pattern 
Type A probably occurred in situ as a 
result of environmental selection pres- 
sures. The present distribution of fes- 
tiva in Middle America is limited to 
lowland forests. Expansion of its range 
in northern parts of the region has been 



prevented by lack of suitable liabitat 
and possibly by competition from Amei- 
va undtdata and Cnemidophorus. Such 
expansion in South America has prob- 
ably been limited by the lack of suit- 
able habitat in Venezuela and eastern 
Colombia and possibly by competition 
from Ameiva hridgesi in western Co- 
lombia. 

The early history of the Ameiva un- 
didata-q uadrdineata-chaitzami complex 
in Middle America may have been simi- 
lar to that of A. festiva. Having origi- 
nated from an unknown South Ameri- 
can ancestral stock, the forerunners of 
the extant species moved into Middle 
America and along both Atlantic and 
Pacific routes. Extensive northward 
movement along the Pacific lowlands 
was probably prohibited by ecological 
barriers, possil^ly the climatic filter bar- 
rier illustrated by Savage ( 1966, Fig. 
23). Northward migration along the 
Atlantic slopes progressed to Nuclear 
Central America and thence, along two 
routes, on into what is now Mexico. 
Presumably, ecological divergence in 
Costa Rica and adjacent Panama, asso- 
ciated with exploitation of beach and 
open-ground habitats, resulted in the 
southern segment of the basal stock be- 
coming A. quadrdineata. The range of 
this species is now disjunct (Fig. 13), 
but in the past it probably extended 
along the Atlantic and Pacific slopes of 
the Western Sierra ( terminology of Ben- 
nett, 1968) in Panama and was joined 
in the lowlands of the isthmus. This 
contention is supported by the simi- 
larity of samples from opposite slopes 
in the eastern part of the ]3resent range 
of the species, a similarity not found be- 
tween samples from opposite slopes 
farther west. Populations of quadrdi- 
neata on Isla del Cano, Costa Rica, and 
Isla de Cebaco and Isla Escudo de 
Veraguas, Panama, apparently owe their 
presence to jiast land connections with 
the mainland during the Pleistocene 
(Bennett, 1968). 

Ameiva widulata, which continued 



70 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



to inhabit more forested areas than 
quadrilineata, migrated northward to 
fill its present range (Fig. 17). Smith 
and Laufe (1946), having to rely on 
Schnchert ( 1935 ) for geological back- 
ground, discussed the subsequent his- 
tory of the species. They postulated a 
broad distribution of tinchilata in Mid- 
dle America during the Miocene, with 
four secondary centers of dispersal, two 
on either side of the Isthmus of Tehuan- 
tepec ( Smith and Laufe, 1946, Fig. 4A ) . 
Further differentiation was promoted by 
the opening of the Tehuantepec portal 
during the Upper Miocene and Lower 
Pliocene and by subsequent migration, 
isolation or ecological divergence of 
populations. If we accept the South 
American origin of the family Teiidae 
(Dunn, 1931), and the timing of the 
reconnection of Middle and South 
America at the Isthmus of Panama as 
Early Pliocene (Lloyd, 1963; Maldo- 
nado-Koerdell, 1964), we must reject 
the hypothesis of Smith and Laufe and 
can eliminate from consideration the 
presence or absence of an open portal at 
the Isthmus of Tehuantepec. It seems 
probable that one line of undulata stock 
moved through Nuclear Central Amer- 
ica along the southern lowlands, cross- 
ing to the Atlantic lowlands through the 
Isthmus of Tehuantepec depression. 
Movement along the Pacific slopes was 
limited by xeric conditions to the north. 
Similar conditions in northeastern Mex- 
ico prevented further northward migra- 
tion on the Atlantic slopes. Eastward 
migration along the Atlantic lowlands 
from the Isthmus of Tehuantepec may 
have been blocked by unsuitable eco- 
logical conditions in the form of 
swamps, or by competition from a sec- 
ond line of undulata stock that had 
migrated along the Atlantic lowlands 
of Nuclear Central America to reach 
Guatemala and the Yucatan Peninsula. 
This second line would have given rise 
to the extant populations of undulata on 
Isla de Cozumel off the Yucatan Penin- 
sula and on Islas del Maiz east of Nica- 



ragua when these islands were con- 
nected with the mainland. Migration of 
the ancestral stock southward from Nu- 
clear Central America into what is now 
Costa Rica would account for the pres- 
ent populations along the southern 
Pacific slopes and on the Meseta Cen- 
tral of Costa Rica. I have postulated 
that the first of the two lines gave rise 
to those populations having individuals 
with color patterns emphasizing stripes 
or combinations of stripes and lateral 
bars or blotches. The second line gave 
rise to those subpopulations character- 
ized by having barred flanks. 

Ameiva chaitzami apparently evolved 
rather recently from a striped fonn of 
undulata, possibly because of ecological 
isolation in the arid valley of the Rio 
Grijalva and its tributaries. If further 
samples linking populations of chait- 
zami with undulata are found, it is pos- 
sible that study will show that chaitzami 
is a form of undulata, as inadvertently 
proposed by Smith and Laufe ( 1946 ) 
and that the sympatry of the two seen 
today in Guatemala is the result of sec- 
ondary contact. 

Although I think it unlikely, studies 
of South American Ameiva may show 
that I have underestimated, rather than 
overestimated, the amount of diver- 
gence of the undulata complex from 
other species. If so, this complex, like 
Cnemidophorus, may be found to have 
its center of radiation in northern Mid- 
dle America and may be considered a 
component of the Middle American 
herpetofaunal element ( = "Hanging 
Relicts," Schmidt, 1943; "Autochthonous 
Middle American Element," Stuart, 
1950 ) proposed by Savage ( 1966 ) . Such 
an arrangement would mean that the 
complex was isolated in Middle Amer- 
ica with the opening of the Panamanian 
portal and, with some adjustments in 
timing, would support the hypothesis of 
Smith and Laufe ( 1946 ) concerning 
evolution of Ameiva undulata itself in 
Middle America. 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



71 



SUMMARY 



The genus Ameiva is broadly dis- 
tributed in Middle and South America, 
as well as on islands throughout the 
Caribbean. The genus is represented in 
Middle America by six species: Aiiwiva 
ameiva (Linnaeus), leptophrys (Cope), 
f estiva ( Lichtenstein and Von Mar- 
tens), quadrilineata ( Hallowell ) , tin- 
dulata (Wiegmann) and chaitzami 
Stuart. These species are variously dis- 
tributed throughout the lowland tropics 
of Middle America from Nayarit and 
central Tamaulipas, Mexico, through 
Panama. Ameiva ameiva and f estiva, 
and possibly leptophrys occur in South 
America as well. Insular populations 
are known of all species, except lepto- 
phrys and chaitzami. 

Geographic variation is extensive in 
scutellation and, in some species, color 
pattern. Sexual dimoiphism and onto- 
genetic change in color pattern is pro- 
nounced in some species. Certain char- 
acters, such as total number of supra- 
labials, number of dorsal granules from 
occiput to rump and scales around the 
tail, are sexually dimorphic in some 
species, not in others. On the other 
hand, several characters show no sexual 
dimorphism at all. Ontogenetic changes 
are most striking among males. Males 
of some species undergo marked 
changes in ventral coloration, especially 
in the gular region; these may be asso- 
ciated with reproductive state. Clines 
are apparent in a great many characters 
and, as might be expected in the context 
of the essential linearity of the Middle 
American land mass, most clines nm 
parallel to the coastlines. This is best 
illustrated by undulata, the species 
having the greatest range in Middle 
America and for which the most samples 
were available. In this species, the 
number of granules around the body 
and the number from occiput to rump 
proved the most sensitive indicators of 
differences between adjacent samples. 
These two characters are relatively 



highly correlated. The numbers and 
arrangement of posterior head scales 
and of supraoculars were found to be 
unstable. The former is especially true 
of leptophrys and, to a lesser extent, 
festiva. The condition of a longitudinal- 
ly divided interparietal, occasionally 
used to diagnose some species of Amei- 
va, was found to be of wide occurrence 
in several species. 

In Middle America, the species of 
Ameiva run an ecological gamut from 
the savanna-dwelling ameiva to festiva, 
which inhabits lowland tropical rain 
forest. In the presence of Cnemido- 
phorus, Ameiva tend toward less open 
habitats, although the same species may 
show more latitude in habitat choice if 
Cnemidophorus is absent. As many as 
three species of Ameiva occur sym- 
patrically in lower Central America. 
Competition among them is partially 
averted by partitioning of the available 
habitat. Typically, one species is a 
forest-dweller, another inhabits open 
areas and the third is intermediate. Size 
differences are also important in this 
context, and partitioning by subadults 
may be different than that by adults. 
Little is known of the demography of 
the Middle American Ameiva. All are 
bisexvial. 

The species of Ameiva of Middle 
America have been assigned to three 
groups. These groups cannot be given 
formal taxonomic recognition until their 
relationships with South American spe- 
cies of Ameiva have been ascertained. 
The composition of the groups may 
have to be modified when these rela- 
tionships are better understood. Group 
I contains only Ameiva ameiva. This 
group has diverged considerably from 
Groups II and III, and can be distin- 
guished on the basis of a large number 
of characters of scutellation and color 
pattern. The group is primarily South 
American in distribution. Group II con- 
sists of festiva and leptophrys, both of 



72 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



which have ranges centered in lower 
Central America. These two species 
have in common such characters as a 
high number of femoral pores and a 
tendency for the posterior head scales 
to be disrupted. Group III is made up 
of unchdata, quadrilineota and chait- 
zami and is entirely Middle American 
in distribution. The group is character- 
ized by certain details of scutellation 
and, especially, color pattern. 

The genus Ameiva is thought to 
have originated in South America and 
to have invaded Middle America fol- 
lowing closure of the Panamanian por- 
tal in Early Pliocene. Of the Middle 
American species, Ameiva ameiva now 
occupies a disjunct range in central and 
western Panama and in South America 
east of the Andes. Ameiva festiva oc- 
curs on eastern and western sides of the 
Andes; it is thought that the eastern 
population gave rise to the Middle 
American population. The species pres- 
ently has a continuous range along the 
Atlantic lowlands to Tabasco, Mexico, 
and along Pacific slopes to southeastern 
Costa Rica. Based on present distribu- 
tion, leptophrys may have evolved as an 
insular isolate in the Panamanian portal. 
Ameiva quadriUneata is thought to be a 
lower Central American derivative of 
early iimhdata stock, and chaitzami a 
recently evolved offshoot of a striped 
form of umlulata. Forerunners of un- 
dulata itself are thought to have mi- 
grated northward on either side of Nu- 
clear Central America. Those moving 
along the Pacific routes gave rise to the 
populations in which striped, barred 
and blotched patterns are emphasized. 
Those moving through the Atlantic low- 
lands have given rise to those popula- 
tions in which lateral bars are empha- 
sized. Populations of tindulata from the 
Atlantic lowlands of Mexico between 



central Tamaulipas and Campeche are 
derivatives of the line which moved 
along the Pacific routes and crossed to 
the Atlantic slopes through the depres- 
sion at the Isthmus of Tehuantepec. 
Without exception, insular populations 
of Ameiva in Middle America occur on 
islands of the continental shelf. These 
populations were isolated from main- 
land populations when rising sea levels 
severed connections with the mainland 
during the Pleistocene. 

The genus Ameiva is the most wide- 
ranging of the exclusively terrestrial 
Neotropical lizards. Over its entire 
range, the genus has exploited a broad 
spectrum of available habitats. For 
these reasons, a better understanding of 
the ecological relationships of Ameiva 
may lead to a better understanding of 
tropical vertebrates in general. To date, 
very little ecological research has dealt 
with Ameiva. Interesting problems in- 
clude the relationships of characters of 
scutellation and color pattern to envi- 
ronmental parameters, the cause(s) and 
significance of color changes in adult 
males, and the ecological relationships 
of sympatric species of Ameiva and 
Cnemidophonis, regardless of whether 
or not these two genera are shown to be 
congeners. Taxonomic problems of im- 
portance to complete understanding of 
the Middle American Ameiva include 
elucidation of their relationships with 
South American species (and the rela- 
tionships of the South American species 
to one another), the relationships of 
chaitzami to undtdata, and the status of 
the population of undulata on the Me- 
seta Central of Costa Rica. These are 
but a few suggestions for future lines of 
research. There seem to be sufficient 
problems to keep a number of investi- 
gators occupied for years to come. 



RESUMEN 



El genero Ameiva esta ampliamente 
distribuido tanto por America Media y 
Sur America, como por las islas del 



Caribe. El genero esta representado en 
America Media por seis especies: Am- 
eiva ameiva (Linnaeus), leptophrys 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



73 



(Cope), f estiva ( Lichtenstein y Von 
Martens), qiiodrilineata ( Hallowell ) , y 
chaitzami Stuart. Estas especies se en- 
cuentran variablemente distribuidas al 
traves de las tierras bajas tropicales de 
America Media, desde Nayarit y Ta- 
maulipas central, Mexico, hasta Pana- 
ma. Ameiva amewa y f estiva, y posible- 
mente leptophrys ocurren tambien en 
Sur America. Se conocen poblaciones 
insulares de todas las especies excepto 
de leptophrys y chaitzami. 

La variacion geografica de la escu- 
telacion y, en algunas especies, el patron 
de coloracion es amplia. El dimorfismo 
sexual y el cambio ontogenetico del 
patron de coloracion es pronunciado en 
algunas especies. Ciertos caracteres, 
como el mimero total de supralabiales, 
numero de granulos dorsales, desde el 
occipito a la anca, y las escamas alrede- 
dor de la cola, son sexualmente dimor- 
ficos en algunas especies pero no en 
otras. Por otro lado, algunos caracteres 
no presentan dimorfismo sexual del 
todo. Los cambios ontogeneticos son 
mas sorprendentes en los machos. Los 
machos de algunas especies sufren cam- 
bios marcados en la coloracion ventral, 
aspecialmente en la region gular, y estos 
se pueden asociar con el estado de re- 
produccion. Gradaciones son aparentes 
en muchos caracteres y en su mayoria, 
corren paralelas a las lineas costera, 
como se puede esperar del contexto del 
arreglo linear de la masa terrestre de la 
America Media. Esto puede ser ilus- 
trado mejor con undulata, la especie 
que tiene la extension mas larga por 
America Media y de la cual se tienen 
el mayor numero de muestras. En esta 
especie, el numero de granulos alrede- 
dor del cuerpo, y el numero desde el 
occipito a la anca probaron ser los mas 
sensibles indicadores de diferencias en- 
tre muestras adjacentes. Estos dos car- 
acteres estan relativemente muy corre- 
lacionados. El numero y arreglo de las 
escamas supraoculares y las posteriores 
a la cabeza es inestable. Lo primero es 
especialmente cierto en leptophrys y, 



en menor grado, festiva. La condicion 
de poseer la escama interparietal divi- 
dida longitudinalmente, empleada oca- 
cionalmente en las diagnosis de algunas 
especies de Ameiva, fue encontrado 
siendo de gran ocurrencia en varias 
especies. 

En la America Media, las especies de 
Ameiva pasan por una gama ecologica 
desde la ameiva que vive en las sabanas 
hasta la festiva que habita en las tierras 
bajas del bosque tropical lluvioso. En 
presencia de CnemidopJiorus, Ameiva 
tiende a vivir en habitats menos abier- 
tos, aunque la misma especie puede 
presentar mayor latitud en la escogen- 
cia del habitat si Cnemidophorus esta 
ausente. Hasta tres especies de Ameiva 
ocurren simpatricamente en la baja Cen- 
tro America. La competencia entre ellas 
se previene por medio de repartimiento 
del habitat utilizable. Tipicamente, una 
especie habita el bosque, otra habita las 
areas/ abiertas y la tercera es inter- 
media. Las diferencias en tamano tam- 
bien son importantes en este contexto, 
y la reparticion del habitat por los sub- 
adultos puede ser diferente de la de los 
adultos. Se sabe muy poco sob re la 
demografia de Ameiva de America Me- 
dia. Todas son bisexuales. 

Las especies de Ameiva de America 
Media han sido asignadas a tres grupos. 
A estos grupos no se les puede dar re- 
conocimiento taxonomico formal hasta 
que sus relaciones con las especies de 
Ameiva de Sur America se determinen. 
La composicion de los grupos podra 
tener que modificarse cuando estas rela- 
ciones se comprendan mejor. El grupo 
I contiene solo a Ameiva ameiva. Este 
grupo se ha apartado considerablemente 
de los grupos II y III, ye se puede dis- 
tinguir en base al gran numero de 
caracteristicas en la escutelacion y pa- 
tron de coloracion. Este grupo es prin- 
cipalmente suramericano en distribu- 
cion. El grupo II consiste de festiva y 
leptophrys, ambas con distribuciones 
concentradas en la baja Centro Amer- 
ica. Estas dos especies tienen en comun 



74 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



caracteres como un numero grande de 
poros femorales y la tendencia a poseer 
escamas rotas en la parte posterior de 
la cabeza. El grupo III se compone de 
undulata, qiiadrilineata y chaitzami y 
esta distribuido enteramente por Amer- 
ica Media. Este grupo esta caracteri- 
zado por ciertos detalles de escutelacion 
y, especialmente, el patron de colora- 



cion. 



Se cree que el genero Ameiva se 
origino en Sur America y que invadio 
la Ameria Media despues del cierre del 
paso panamefio en el Plioceno tempra- 
no. Ameiva ameiva posee una distribu- 
cion discontinua en Panama central y 
oeste, y en Sur America al este de los 
Andes. Ameiva f estiva ocurre en los 
lados este y oeste do los Andes; se cree 
que la poblacion del oeste dio origen a 
la poblacion de America Media. Esta 
especie ahora tiene una distribucion 
continua a lo largo de las tierras bajas 
del Atlantico hasta Tabasco, Mexico, y 
a lo largo del Pacifico hasta el sureste 
de Costa Rica. Basado en la distribu- 
cion presente, leptophrys pudo haber 
evolucionado de un aislamiento insular 
en el paso panamefio. Se cree que quad- 
rilineata se derivo, en la parte baja de 
Centro America, de una linea temprana 
de undulata y chaitzami evoliciono 
recientemente de una forma de undu- 
lata listada. Se cree que los precursores 
de undulata misma emigraron hacia al 
norte por ambos lados de Centro Amer- 
ica unclear. Esos que tomaron las rutas 
del lado del Pacifico dieron origen a las 
poblaciones en las cuales las listas, 
barras, y manchas son enfatizadas. Las 
que se desplazaron por las tierras bajas 
del Atlantico dieron origen a las pobla- 
ciones en las cuales se enfatizan las 
barras laterales. Las poblaciones de 
undulata de las tierras bajas del Atlan- 
tico de Mexico, entre Tamaulipas cen- 
tral y Campeche, son derivadas de la 
linea que se movio a lo largo de las 



rutas del Pacifico y que crusaron la de- 
presion del istmo de Tehuantepec para 
dirigirse a la region Atlantica. Sin ex- 
cepcion, las poblaciones insulares de 
Ameiva en America Media ocurren en 
islas de la plataforma continental. Estas 
poblaciones fueron aisladas de las pob- 
laciones del continente cuando los ni- 
veles del mar subieron durante el Pleis- 
toceno y separaron las conecciones con 
tierra firme. 

El genero Ameiva es el mas amplia- 
mente distribuido de las lagartijas Neo- 
tropicales terrestres. Sobre su total dis- 
tribucion, el genero ha expotado un 
gran espectro de habitats utilizables. 
Por estas razones, una mejor compren- 
sion de las relaciones ecologicas en 
Ameiva puede llevar a una mejor com- 
prension de los vertebrados tropicales 
en general. Hasta esta fecha, muy poca 
investigacion ecologia a tratado sobre 
Ameiva. Problemas interresantes inclu- 
yen las relaciones de los caracteres de 
escutelacion y el patron de coloracion 
con parametros ambientales, la causa(s) 
y significado de los combios de colora- 
cion en machos adultos, y las relaciones 
ecologicas entre especies simpatricas de 
Ameiva y Cnemidophorus, indiferente- 
mente de presentar o no a los dos ge- 
neros como si fuesen congenericos. 
Problemas taxonomicos de importancia 
para completar la comprension de Am,- 
eiva de America Media incluyen la 
aclaracion de sus relaciones con las 
especies de Sur America (y las rela- 
ciones de las especies suramericanas 
entre si), las relaciones entre chaitzami 
y undulata, y la posicion relativa o 
status de undulata en la Meseta Central 
de Costa Rica. Estas son no mas que 
algunas sugerencias para futuras lineas 
de investigacion. Parece haber sufici- 
entes problemas para mantener ocupa- 
dos a un numero de investigadores 
durante varios aiios. 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



75 



APPENDIX A 

An example of summarized data as used in this report, along with results of 
Gabriel's Simultaneous Test Procedure analysis for number of femoral pores of Pana- 
manian Ameiva amewa. For a complete set of the statistical results of this study, 
order NAPS Document #01660 from CCM Information Corporation, 866 3rd Ave- 
nue, New York, New York 10022; remitting $2.00 for microfiche or $7.00 for photo- 
copies. 



Ameiva amewa: Femoral Pores 



Sample 
Number 


X 


Standard 
Deviation 


Sample 
Size 


Range 


STP 


6 


35.0 


2.76 


23 


29-40 




5 


34.1 


1.64 


11 


31-36 






7 


32.4 


1.85 


20 


29-37 








3 


32.3 


2.07 


24 


28-36 








2 


31.8 


1.93 


21 


29-35 








1 


31.7 


2.61 


40 


25-38 








4 


30.1 


1.28 


13 


27-32 






All 


32.4 


2.69 


270 


25-41 







76 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



APPENDIX B 

Specimens Examined 



Ameiva ameiva 

Sample 1. PANAMA: Canal Zone: An- 
con, MCZ 19805-11, UMMZ 98415-16, 98418; 
Balboa, AMNH 37129, FMNH 13441, UMMZ 
98411-14; Bruja Point, FMNH 13445; nr 
Corozal, MCZ 19813; Diablo Heights, UMMZ 
98417; 6.4 km E Lacone, KU 62701; Mira- 
flores, FMNH 16584-5. Panama: nr City of 
Panama, MCZ 9924-27, 9929-30, 9933-41; 
Panama City, El Cangrejo, KU 96742-43; 
Tapia, AMNH 28005-6, 28008. 

Sample 2. PANAMA: Canal Zone: Gatiin, 
FMNH 16555-57, 16572-74, 16580, 16582-83; 
Cristobal, FMNH 34163-64. Colon: Achiote, 
KU 76109-18, 96737. 

Sam^ple 3. PANAMA: Panama: Bejuco, 
KU 107546; Nuevo Gorgona, AMNH 89925- 
46; ca 12 km NW San Carlos on road to El 
Valle, KU 107547. 

Sample 4. PANAMA: Panama: Archi- 
pelago de las Perlas, Isla San Jose, AMNH 
66491, USNM 120611-23. 

Sample 5. PANAMA: Herrera: Las Minas, 
ANSP 22452; Parita, USNM 127291-94, 
127296; 8 km SE Parita, KU 107538-39; Pese, 
KU 107540-45. 

Sample 6. PANAMA: Los Santos: Gua- 
nico, KU 107521-35; Playa Guanico, near 
mouth of Rio Guanico, KU 107909; Las Pal- 
mitas, USNM 148207-11, KU 110731-34. 

Sample 7. PANAMA: Panama: S Slope 
Cerro Campana, KU 76094-106, 107548-54. 

Additional Specimens Not Used In STP 
Analysis. COSTA RICA: No definite locality, 
BMNH 80.6.21.1-2. PANAMA: Canal Zone: 
Alajuela, UMMZ 76104(4); Ancon, KU 
110740; Bas Obispo, USNM 54129; Camp 
Chagres, KU 76107-8; Chagres, USNM 5519; 
Culebra, FMNH 16559-61; Culebra Cut, 
FMNH 16554; Fort Kobbe, KU 110739; Fort 
Kobbe, Venado River, KU 110742-43; El 
Vigia, UMMZ 76012; Frijoles, UMMZ 63760; 
Gamboa, AMNH 32818; Juan Mina, Chagres 
River, KU 108258-61; Lower Chagres, AMNH 
37073-4, 37077; Madden Dam, UMMZ 
76015(2); Madden Forest Preserve, KU 
96740-41; Rio Abajo, FMNH 130685; Rio 
Frijoles, USNM 8394; Road K9, FMNH 
130972; Rosseau, KU 62702; Summit, FMNH 
130683, 131309; nr Summit Experimental 
Gardens, UMMZ 95484; Tabernilla, FMNH 
16585-87, USNM 54130. Chiriqui: 2.5 km E 
La Concepcion, KU 96736; Remedios, AMNH 
28390; 3 km W Rio Chorcha, KU 95540-44; 
16 km W Rio Chorcha, KU 95545; Rio Salado, 
KU 95546-47. Cocle: Bank of Rio Chico de 
Nata, KU 95548-52; El Valle, AMNH 89924; 
16 km S, 9 km W Penonome, KU 107536; 19 
km S, 9 km W Penonome, Puerto Posoda, KU 



107537. Colon: Portobello, USNM 65122; 
Quebrada Bonita, KU 96738-39; Santa Rosa, 
AMNH 71705. Herrera: Tres Puntas, ANSP 
22470-74. Los Santos: Jobero, KU 110735- 
36; Pocri, KU 107520; Punta Mala, AMNH 
71704. Panama: Cafiitas, UMMZ 124148-50; 
Cerro Azul, AMNH 89947-48; Chorrera, 
AMNH 71703, FMNH 57597-98; Flora de 
Lagunas (Rio Caiiitas), UMMZ 124146-47; 
Gatuncillo, UMMZ 76013; Isla San Miguel, 
MCZ 9942-47; Isla Taboga, FMNH 16576, 
USNM 54197, 54205, 102754-56; Isla Tabo- 
guilla, USNM 102809-10; Juan Diaz, KU 
110737-38; Nuevo Emperador, FMNH 130937, 
130939-45, KU 110471; Pacora, AMNH 32817, 
FMNH 120956; Rio Bayano, USNM 53980; 
Rio Mamoni, FMNH 16566-68; Rio Mamoni, 
5 km E Chepo, KU 76119-25; Rio Tapia, 24 
km E Panama City, AMNH 42953. Veragita^-: 
26 km W Santiago, KU 95553-55; Sona, 
USNM 51972-73. 

Ameiva chaitzami 

Sample 1. MEXICO: Chiapas: Comitan, 
UMMZ 94905(24), 94906(5), 94907. 

Sample 2. GUATEMALA: Htiehuete- 
nango: San Antonio Huista, UMMZ 120192; 
1 km N San Antonio Huista, UMMZ 120189, 
120190(4), 120191(2); 1 km W San Antonio 
Huista, UMMZ 120195; 1-2 km W San An- 
tonio Huista, UMMZ 120193(18); 1-2 km E 
San Antonio Huista, UMMZ 120194(4). 

Sainple 3. GUATEMALA: Alta Verapaz: 
Languin-Cahabon road, MCZ 52170; On Lan- 
guin-Cahal^on road ca 2 km N Finca Canihor, 
UMMZ 90638-41, 90643. El Peten: 1 km N 
Popti'm, UMMZ 124355; 2 km SW Poptun, 
UMMZ 124356; 2 km NE Poptun nr source 
Rio San Juan, UMMZ 124357. 

Additional Specimens Not Used In STP 
Analysis. MEXICO: Chiapas: La Libertad, 
MCZ 56007, USNM 137216; La Libertad, nr 
Rio Cuilco where it crosses the Guatemalan 
border, FMNH 100006, 100712-15; unknown, 
but pro1ia]:)ly near La Libertad, FMNH 
100716. 

Ameiva festiva 

Sample 1. GUATEMALA: El Peten: 
Piedras Negras, FMNH 115429-43, USNM 
133826-36, UIMNH 11349-62. 

Sample 2. GUATEMALA: Alta Verapaz: 
Chinaja, KU 55839, 55849-50, 59566; ca 3 km 
S Chinaja, KU 55841-48, 55857. El Peten: 6 
km NNW Chinaja, KU 55836; 11 km NNW 
Chinaja, KU 55837-38; ca 15 km NW Chinaja, 
KU 55851-56, 59567-73; Toocog, 15 km SE 
La Libertad, KU 55840. 

Sample 3. GUATEMALA: Alta Verapaz: 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



77 



Finca Chama, UMMZ 91319, 91321(2), 
91322(2), 91323(3), 91324(2), 91325(2), 
91326(4), 91327, 91328(3), 91329(2), 91330, 
91331(8), 91332. 

Saiyiple 4. HONDURAS: Atlantida: ca 
15 km E La Ceiba, mtns above Corozal, JRM 
2299, 2418-21, 2439-40, 2466-69; Along Rio 
Cangrejal ca 12 km SSE La Ceiba, KU 
101195-97; Tela, Lancetilla, UMMZ 70324(3). 

Sample 5. NICARAGUA: Zelaya: Bo- 
nanza, KU 84870-72, 86039-40, 101201-12; 6 
km E Bonanza, KU 101198; On banks of Rio 
Tunkey 4 km E Bonanza, KU 101199-200; 
Eden Mine, ANSP 2133-35. 

Sample 6. NICARAGUA: Zelaya: Cukra, 
AMNH 16587-96; Kanawa, AMNH 16597, 
16605-16; Between Cukra and Kanawa, 
AMNH 16600, 16602-4. 

Sample 7. NICARAGUA: Zelaya: El 
Recreo, 16 km W Rama on Rio Mico, UCLA 
9769-70; El Recreo, S side Rio Mico, KU 
101874; 16 km above El Recreo on Rio Mico, 
UMMZ 79838(12); 11.2 km above Rama, 
UMMZ 79840(5), 79841(2); Junctions Rio 
Mico and Rio Siquia, UMMZ 79839(4). 

Satnple 8. NICARAGUA: Rio San Juan: 
Tuli Creek, AMNH 16666-74, 16676-700. 

Sample 9. COSTA RICA: Heredia: Puer- 
to Viejo, KU 67298-309, 91787, 104081-82; 
UMMZ 125505; 1.5 km N Puerto Viejo, KU 
67310-11; 2 km NE Puerto Viejo, KU 95558- 
68; 3 km S Puerto Viejo, KU 95556-57; ca 
12.8 km NE Puerto Viejo along Rio Sarapiqui, 
UMMZ 125506(2); Rio Sarapiqui at Puerto 
Viejo, UMMZ 123582-83; Finca La Selva on 
Rio Puerto Viejo, KU 100683-85. 

Sample 10. COSTA RICA: Cartago: Tur- 
rialba, FMNH 101906, 103107, MCZ 55091- 
92, 55094, USNM 37737, KU 25633-34, 
25636-39, 34783-89, 34826-27, 40552, UMMZ 
117492; 5 km SE Turrialba, KU 25174-75; 
6.4 km SE Turrialba, KU 34778-79; ca 4 km 
SW Moravia de Turrialba, KU 67312; 4.8 km 
W Turrialba, Hacienda Florencia, KU 34790- 
91. 

Sajnple 11. COSTA RICA: Limon: Vi- 
cinity of mouth of Rio Tortuguero, AMNH 
89189-203, 89205-15. 

Sample 12. COSTA RICA: Limon: Gua- 
piles, MCZ 15335-36; La Lola, KU 34798-801, 
34807-13, UMMZ 117493(9); Los Diamantes, 
FMNH 101250, 101252-54, 101256-59, 
101261-63, 101904-5, 101907, KU 25581-86, 
25619-32, 34772-76, 34802-6; Puerto Limon, 
KU 34797; Zent, MCZ 15376-81. 

Sample 13. PANAMA: Bocas del Tow: 
Almirante, FMNH 154445-48, 154451, MCZ 
19785-87, KU 80207-11; 3.2 km NW Almi- 
rante, KU 96745-46; 4.8 km NW Almirante, 
KU 96747; 5.6 km NW Almirante, UU 5162; 
9.6 km NW Almirante, KU 96748; 12.8 km 
NW Almirante, KU 96749-52. 

Satnple 14. PANAMA: Bocas del Tow: 



Isla Escudo de Veraguas, KU 108262-71, UU 
5153-54. 

Sample 15. PANAMA: Canal Zone: Bar- 
ro Colorado Island, AMNH 6.3401, 89949, 
FMNH 13342-43, MCZ 20612-13, 28066-67, 
KU 76126-27, UMMZ 63746-55, 63766, 63779, 
124156. 

Sample 16. PANAMA: Darien: Rio 
Mono, ca 5 km above junction with Rio Tuira, 
KU 96781-84; Rio Tuira at Rio Mono, KU 
96785-804. 

Sample 17. COSTA RICA: Puntarenas: 
Agua Buena, Caiias Gordas, KU 40545-48, 
40553-58; Golfito, KU 67313; 2.5 km ESE 
Piedras Blancas, KU 93978; Rio Zapote, 8 km 
E Palmar Norte, KU 93977, 95569. 

Sample 18. COSTA RICA: Puntarenas: 
3 km NW Rincon de Osa, KU 101487-88; 7.2 
km SE Rincon de Osa, KU 101489-90; 9.6 
km NW Rincon de Osa, KU 101483-86; 10 km 
SSE Rincon de Osa, KU 101473-80; 11.7 km 
SSE Rincon de Osa, KU 101481-82. 

Sample 19. COLOMBIA: Cordoba: Cafio 
Guarmal, FMNH 165260-66; Upper Rio Ure, 
FMNH 165225, 165227-30, 165748-67, 
165769. 

Sainple 20. COLOMBIA: Choco: An- 
dagoya, USNM 124246; Andagoya-Condoto 
Area, UMMZ 121463; Condoto, UMMZ 
121465(13); Tado, ANSP 26265-66. 

Additional Specimens Not Used In STP 
Analysis. BRITISH HONDURAS: Cayo: 
Valentin, UMMZ 92372; Stann Creek: Mid- 
dlesex, FMNH 4460; Toledo: 1.6 km E Swa- 
zey Branch of Monkey River, MCZ 71606-7; 
District Unknown: Silk Grass Creek State For- 
est, FMNH 4459; Bobowina, FMNH 49313-14. 
COLOMBIA: Boyaca: Muzo, MCZ 56233; 
Choco: Boca de la Raspadura, AMNH 18266- 
68; El Valle, Bahia Solano, Mutis, USNM 
151512; El Valle, Pizarro, USNM 151513-14; 
Pizarro, FMNH 43816-28; Rio San Pado, 1.6 hn 
below Boca de la Raspadura, AMNH 18274; Si- 
erra Bauda, ANSP 25526; Sierra de Darien, 
ANSP 25527-28; Upper Rio San Juan, FMNH 
15652-53; Santander: El Centro Santander, 
ANSP 25075, 25427; Departamento Unknown: 
E base Cordillera de Bogota at extreme limit 
of llanos, ANSP 24165; "Tambo S. Monica, 
Atlantic Slope," AMNH 18265; Region of 
Truando, ANSP 9071. COSTA RICA: Ala- 
juela: 3.2 km NE Muelle Canal, UCR 23-26; 
Isla Bonita, FMNH 103103-6; Cartago: nr 
Peralta, Lake Bonilla, KU 43886-8; nr Peralta, 
Tunnel Camp, KU 34828, 38825; Reventazon, 
USNM 37492; Rio Pacuare, nr Pacuare, KU 
34777; Suiza, FMNH 101251, 101255, KU 
25635, 34792; Turrialba, FMNH 101909; 6.4- 
8.0 km S Turrialba, FMNH 101260; Guana- 
caste: Laguna de Arenal, UCR 232-33, 268- 
69; Tenorio, KU 34814-24; Tilaran, ANSP 
24561; 5 km ENE Tilaran, KU 40551; Tron- 
cadero, nr Lake Arenal, KU 40539-40; Here- 



78 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



dia: Barro Colorado, KU 100679-80; Behveen 
Puerto Viejo and Cariblanco, KU 40537-38; 
Limon: Bataan, KU 34780-82; Pandora, KU 
86573-76, 100681-82; Suretka, MCZ 19777, 
KU 40541-44, 40559-63; Puntarenas: Agua 
Buena, Caiias Gordas, KU 40549-50; San Jose: 
San Isidro del General, FMNH 101908, 
101910, KU 34793-94; 8 km E San Isidro del 
General, FMNH 120236; 24-32 km NW San 
Isidro del General, San Juan Mtns., KU 34795- 
96; Province Unknown: Guayabo, FMNH 
6167; Rio Frio, USNM 19534-36; Locality 
Unknown, USNM 30558, 32612-13. GUATE- 
MALA: Alfa Verapaz: Cubiliquitz, UMMZ 
91334, 91335(2), 91336(2); Finca Chama, 
MCZ 28452-54, UMMZ 91320(2), 91329, 
91331; Panzos, UMMZ 9137; Samazana, 
UMMZ 91333(4); El Quiche: Finca Tesoro, 
UMMZ 89197(4); Izabdh "Babos Pi., Mt. 
Playita," FMNH 20081, 20109-11; Quiriqua, 
FMNH 20487-88; El Peten: Pacomon, USNM 
71394; Poptim-San Luis Road, ca 15 km S 
Poptim, UMMZ 124359; 12 km S PopUm, 
UMMZ 124360; 10 km NW Poptun, UMMZ 
124.361(5); Ramate, USNM 71406, 71817, 
71823, 71826-27, 71829-31, 71836; Localiiij 
Unknown: MNHN 5480. HONDURAS: Co- 
pan: 19.2 km ENE Copan, JRM 2895, 2897; 
Cortes: Hacienda Santa Ana, San Pedro Sula, 
FMNH 5067-71; Mtns W of San Pedro Sula, 
FMNH 5074; Garcias a Dios: Rio Segovia, 
USNM 24527-28; Santa Barbara: Jaral, FMNH 
5073; Yoro: Mataderos Mountains, FMNH 
21781, MCZ 38924; Portillo Grande, MCZ 
38925; Subirana Valley, MCZ 38926; De- 
partamento Unknown: Carmelina, USNM 
62972; Lake Ticamaya, FMNH 5072. MEX- 
ICO: Chiapas: Palenque, USNM 133818-25, 
UIMNH 11343, KU 94107-8; Tabasco: Teapa, 
USNM 47454-56. NICARAGUA: Rio San 
Juan: Greytown, USNM 19533-34, 19640-1; 
Zelatfa: 3-4 km W Bluefields, KU 101213-15; 
Cupitna, AMNH 16633-51; Rio Escondido 70 
km from Bluefields, USNM 19735, 19871. 
PANAMA: Boca,s- del Toro: 11 km NW Al- 
mirante, FMNH 68171; Bastimentos, USNM 
150021; Boca del Drago, USNM 142332-33; 
Cayo Agua, USNM 150014-15; Rio Cahuita, 
ca 5 km above mouth, KU 108278-79; Rio 
Changena, FMNH 130971, KU 110744-45; 
Canal Zone: Cerro Azul, FMNH 130684; 
Cristobal, FMNH 34165; Gamboa, FMNH 
131308; Gatun, FMNH 16654-56, USNM 
54134; Chiriqui: Boquete, UMMZ 58296; El 
Volcan, USNM 129924; Finca Palosanto 7 km 
NW El Volcan, Rio Chiriqui Viejo, KU 96808- 
11; Finca Santa Clara, KU 110753-54; Volcan 
Chiriqui, UMMZ 101787; Code: El Valle, 
FMNH 47456; Finca El Valle de Anton, 
FMNH 60174; CoUn: Achiote, KU 76129-30; 
Candelaria Hydrographic Station, FMNH 
68172-73; Portobello, FMNH 16652-53, USNM 
54072-73, 80044; 3.5 km SE Puerto Pilon, 



KU 110755; Rio Candelaria nr Hydrographic 
Station, FMNH 68174-76; Darien: Cafia, 
USNM 50134, 50136, 50138-39, ,50142-43; El 
Real, KU 80529; Laguna, KU 76131; Rio 
Cupe, ca 12 km SSW Boca de Cupe, KU 
96805; Rio Esnape, Sambu Valley, MCZ 
17224-29; Rio Jaque, 1.5 km above Rio 
Imamado, KU 110751-52; Rio Jesuito, 
Sapo Mtns, MCZ 17179-80, 17189; Rio 
San Antonio nr base of La Jarcia, KU 110750; 
Tacarcuna, KU 76132-39; Tacarcuna Village, 
USNM 141819-20; Panama: S Slope Cerro 
Campana, KU 76128; Cerro Tefe, KU 96806-7; 
E Slope Cerro Jefe, KU 80530-37; NW Slope 
Cerro Prominente, KU 80538-39; Chepo, 
FMNH 16656; El Valle de Anton, trail to La 
Mesa, FMNH 68177-79; Rio Abajo, FMNH 
57599; Rio Siluganti, UMMZ 124151-55; San 
Bias: Camp Sasardi, KU 110748-49; Camp 
Summit, KU 110746-47; Veraguas: Mouth of 
Rio Concepcion, KU 108272-77; Province Uti- 
known: Cabima, USNM 48504; Chico, UMMZ 
76011; Nombre de Dios, MCZ 26944; Punta 
de Pena, USNM 38693-98; "Veragua," ZMB 
881a, 881b. 

Ameiva leptophrys 

Sample 1. COSTA RICA; Puntarenas: 
Camp Seattle, 6.4 km inland from Rincon de 
Osa, USC(CRE) 752; Vicinity of Rincon de 
Osa, KU 101491-528, 101825. 

Sample 2. COSTA RICA: Puntarenas: 
Goto, Colfo Dulce, UMMZ 71999-72002; Es- 
quinas Forest Preserve between Golfito and 
Palmar, KU 34766-70; 3.2-4.8 km W Palmar 
Norte on road to Puerto Cortez, USC(CRE) 
7101c-7101f; 6 km W Palmar Norte, KU 
93982-83, 95570-82; 6 km SE Palmar Norte, 
KU 67314-16; Rio Zapote, 8 km E Palmar 
Norte, KU 93980-81; Villa Neilly, nr town of 
Corredor, USC(CRE) 179a; 12.3 km WNW 
\'il]a Neilly, KU 67317. 

Sample 3. PANAMA: Canal Zone: Barro 
Colorado Lsland, AMNH 47018-19, 63403(2), 
89953, ANSP 23021-22, FMNH 13307, 1.3344- 
47, 13375-76, MCZ 28063-65, 29386, KU 
76140-45, 80540, UMMZ 63756-59, 63780; nr 
Coro/al, MCZ 19814; Corozal Hill, USNM 
102743. 

Sample 4. PANAMA: Darien: Rio Chu- 
cunaque, ca 7 km above Rio Morti, KU 
107561-63; Rio Cupe, ca 12 km SSW Boca de 
Cupe, KU 96815; Rio Tuira at Rio Mono, KU 
96816-25; Rio Ucurganti, ca 7 km above 
mouth, KU 107564-65; Tacarcuna, KU 76158- 
61; Tacarcuna Village, USNM 141817-18; 
Three-Falls Creek, AMNH 37870. 

Additional Specimens Not Used In STP 
Anahisis. COSTA RIGA: Puntarenas: Par- 
rita. La Tulieta, Unca La Ligia, USC(CRE) 
8256, 8260]>8260c; 2.5 km ESE Piedras Blan- 
cas, KU 93979; Rincon de Osa, RDS 1122-23, 
1170, 1177; San Jose: San Isidro del General, 
FMNH 103111, 103117, KU 25640-42; Prov- 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



79 



ince Unknown: Pozo Azul de Parris, ANSP 
21470; Santo Domingo de San Mateo, USNM 
37754. PANAMA: Canal Zone: Alajuela, 
UMMZ 76009; Camp Chagres, KU 76146-48; 
Corozal, AMNH 67073; Gatim, USNM 54142; 
0.5 km S Juan Mina, FMNH 68159; Madden 
Dam, UMMZ 76010; Miraflores, FMNH 16650; 
Rio Cardenas, Corozal, USNM 53868; Chiri- 
qui: Progreso, MCZ 19865; 13 km N Puerto 
Armuelles, KU 96814; Code: El Valle, AMNH 
89950-52, USNM 129906, KU 76162-63; 
Colon: Achiote, KU 76149-51; Portobello, 
USNM 92593; Quebrado Bonita nr Buenavista, 
KU 96812; 1 km W Rio Piedras, KU 96813; 
San Bias Point, AMNH 71706; Darien: Cana, 
USNM 50135, 50137, 50140-41; Canclones, 
UMMZ 124975; nr mouth of Rio Canclones, 
UMMZ 124974; Taque, KU 110767-68; La- 
guna, KU 76156-57; Mt. Sapo, ANSP 22963, 
MCZ 17201-3; Rio Jaque, 1.5 km above Rio 
Imamodo, KU 110769-70; Rio Jesuito, Sapo 
Mtns, MCZ 17196-97, 17199; Herrera: Las 
Minas, ANSP 22450; Los Santos: N Slope 
Cerro Cambutal, KU 107555-56; Guanico 
Abajo, KU 110771; Panama: Cerro Azul, 
USNM 54136, 54147; Cliffs W of Cerro Cam- 
pana, FMNH 60178-79; S Slope Cerro Cam- 
pana, KU 76152-53, 107558-59; Chepo, FMNH 
16645; Las Cumbres, KU 96827; Nuevo Em- 
perador, FMNH 130938; nr Panama City, 
MCZ 10925, 10927; 2.8 km ENE Panama 
City, Rio Pacora, KU 96826; 9 km NNE Pa- 
cora, Rio Pacora, KU 107557; Rio Mamoni, 5 
km E Chepo, KU 76154-55; Rio Siluganti, 
UMMZ 124144-45; Rio Tocumen, MCZ 19816; 
San Bias: Camp Sasardi, KU 110756-65; Camp 
Summit, KU 110766; Veragtias: Mangillo, 
ANSP 22450. 

Ameiva quadrilineata 

Sample 1. NICARAGUA: Rio San Juan: 
Greytown, USNM 19537-45, 20741-42, 32230; 
Rio San Juan, USNM 24984. 

Sample 2. COSTA RICA: Limon: Colo- 
rado Bar, AMNH 16726-28; Tortuguero, 
USC(CRE) 2653(2); Vicinity of mouth of 
Rio Tortuguero, AMNH 89217-46. 

Sample 3. COSTA RICA: Liinon: Los 
Diamantes, UMMZ 117491; La Lola, USC 
(CRE) 128(3), 8063, 8070(3); Zent, MCZ 
15382-85. 

Sample 4. PANAMA: Bocas del Tow: 
Almirante, FMNH 60171-72; Careening Cay, 
KU 96840-44; Cayo Zapatillo Grande, KU 
96845-51; Isla Bastimentos, Bastimentos, KU 
96831-39; Isla Bastimentos, Punta Vieja, 
USNM 150016-20, 150022-23; Isla Colon, 
road from Bocas to la Gruta, KU 96829-30. 

Sample 5. PANAMA: Bocas- del Tow: 
Isla Escudo de Veraguas, KU 108280-85, UU 
5166. 

Sample 6. COSTA RICA: San Jose: San 
Isidro del General, FMNH 103115, KU 25643- 



50, 34685-89, USC(CRE) 2742; 4.8 km NE 
San Isidro del General, USC (CRE) 2843; 5.4 
km SW San Isidro del General, UMMZ 
117490(2); 3.2 km W San Isidro del Gen- 
eral, KU 25651; 8 km SE San Isidro del 
General, KU 95583-90. 

Sample 7. COSTA RICA: Ptmtarenas: 
1.5 km E Palmar Norte, KU 95592-93; 6 km 
W Palmar Norte, KU 95633-40; 3.2-4.8 km W 
Palmar Norte on road to Puerto Cortez, USC 
(CRE) 7101a-7101b; 5 km SE Palmar Sur, 
KU 67318-19; 6 hn SE Palmar Sur, KU 
67320-25; Rio Zapote, 8 km E Palmar Norte, 
KU 95607-20, 95622-26, 104083-85. 

Sample 8. COSTA RICA: Puntarenas: 
Rincon de Osa, RDS 1136-38, 1144, 1147, 
1152, 1154-63, KU 101463-69, USC (CRE) 
7239(14); 3 km NW Rincon de Osa, KU 
101470-72; 11.7 km SSE Rincon de Osa, KU 
101459-62. 

Sample 9. PANAMA: Chiriqui: Comarca 
del Baru, Puerto Annuelles, FMNH 68166-67, 
Progreso, UMMZ 58186-202; 8 km S Progreso, 
KU 96852; Puerto Annuelles, MCZ 45668-71; 
13 km N Puerto Annuelles, KU 96853-65. 

Sample 10. PANAMA: Chiriqui: Bo- 
quete, ANSP 21941-43, UMMZ 58183-85, 
58209-18. 

Additional Specimens Not Used In STP 
Analysis. COSTA RICA: Li7n6n: Colorado 
Bar, AMNH 16729-75; La ForUina, AMNH 
99679; Old Harbor, USNM 32614-16; Suretka, 
KU 40578; Puntarenas: 0.8-1.6 km E Bani 
along N bank of Rio Baru, USC (CRE) 7096; 
Buenos Aires, FMNH 2519; Isla del Cano, 
UMMZ 71194; Goto, UMMZ 72005-6; El Gen- 
eral (Viejo) between Palmar Sur and Golfito, 
KU 34690; Esquinas, KU 34691; Golfito, 
USC(CRE) 7111(5), 7233(3); Km 47 on 
railway to Golfito, USC (CRE) 176(8), 177 
(2), 178; Finca El Helechales, 15 km NE 
Potrero Grande, USC (CRE) 8268, 8271; 27 
km E Palmar Norte, Rio Punta Nuevo, KU 
95596-606; 30 km E Palmar Norte, Rio La 
Vieja, KU 95627-32; 2-5 km ESE Piedras 
Blancas, KU 93984-86; Puerto U\dla, UMMZ 
72004; Rincon de Osa, RDS 1139-40, 1142, 
1145-46, 1148-51, 1153; San Isidro del Gen- 
eral, FMNH 103116; 22 km SE San Isidro del 
General, USC (CRE) 2648; 57 km SE San 
Isidro del General, KU 95591; Villa Neilly, 
USC(CRE) 179; 1.6 k-m E Volcan de Buenos 
Aires, Cone Finca, UMMZ 117573; San Jose: 
San Isidro del General, 3.2 km W Intabee, 
FMNH 103102. NICARAGUA: Rio San 
Juan: Greytown, USNM 6053(2). PANAMA: 
Chiriqui: SW Slope Cerro Homito, KU 96867; 
11 km NE Gualaco Road to Valle Homito, KU 
96866; El Hato, edge of lava flow, FMNH 
130686; Llano del Volcan above Paso Ancho, 
FMNH 60175-77, 68168-70; Herrera: Las 
Minas, ANSP 22454-55; Veraguas: 8 km SW 
El Maria, KU 107566; Isla Cebaco, KU 96828. 



80 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



Ameiva undulata 

Sample 1. MEXICO: Tamaulipas: Go- 
mez Farias, UMMZ 110802, 110839; nr Gomez 
Farias, UMMZ 111145; 8 km NE Gomez 
Farias along Rio Sabinas, UMMZ 101476-78, 
101479(6), 101480-82, 101515-16, 104059 
(7), 105489, 111142-44; Pano Ayiictle, 8 km 
NE Gomez Farias, UMMZ 98982. 

Sample 2. MfiXICO: San Luis Potosi: 
Axtla, AMNH 67341; Hiiichihuayan, USNM 
133859; Tamazimchale, AMNH 66065-7, 
FMNH 38611, UIMNH 27360, 51207; 3.2 km 
NE Tamazunchale, TCWC 4090-92, UMMZ 

119819, UIMNH 16804; Xilitla Region, KU 
24056-7. 

Sample 3. MEXICO: Veracruz: Tierra 
Colorado, FMNH 75793, 126933-41, MCZ 
56004. 

Sample 4. MfiXICO: Veracruz: Cuaut- 
lapam, KU 105823, UMMZ 41423-32, 41434- 
42, 41444-51, 41453-54. 

Sample 5. MfiXICO: Veracruz: Cate- 
maco, UMMZ 118727; 0.8 km W Laguna de 
Catemaco, UMMZ 126412; 4 km NE Cate- 
maco, UMMZ 126409; Goyame, UIMNH 
36877, 39217-18, UMMZ 111456, 114794; 1 
km S Dos Amates, between Sontecomapan 
and Catemaco, UMMZ 126413; Laguna En- 
cantada, 3 km NE San Andres Tuxtla, UMMZ 

119820, 126407; Rancho El Tular, ca 8 km N 
San Andres Tuxtla, UIMNH 39212; San An- 
dres Tuxtla, USNM 46899-904, UIMNH 
24691, 27347, 28054, 28056; nr San Andres 
Tuxtla, FMNH 126709; 2 km NE San Andres 
Tuxtla, UMMZ 121156; ca 4 km N San An- 
dres Tuxtla, UIMNH 39212; San Martin 
Mountains, UIMNH 35460; Sontecomapan, 
UMMZ 114795, 126408. 

Sample 6. MfiXICO: Veracruz: Hacienda 
La Oaxaquena, AMNH 62331-33; 20 km S 
Jesus Carranza, KU 23953, 24186; 25 km SE 
Jesus Carranza, KU 26956-57, 26959, 26962- 
63, 26965-71; 35 km SE Jesus Carranza, KU 
23950; 20 km ENE Jesus Carranza, KU 26955, 
26961. 

Sample 7. MfiXICO: Tabasco: Frontera, 
USNM 25091, 46659, 47453; 16 km SW Fron- 
tera, KU 95669-78. 

Sample 8. MfiXICO: Campeche: Bal- 
chacaj, FMNH 106716-18, 106721, UIMNH 
26167-70; Tres Brazos, FMNH 106719, 
UIMNH 26172-75, UMMZ 81924. 

Sample 9. MfiXICO: Campeche: Dzil- 
balchen, KU 75589-98. 

Sample 10. MfiXICO: Yucatan: Piste, 
KU 70598-600, 70604-5, 70609, 70611-15, 
70617, 70620-26, 70628-29, 70631-32, 70634- 
35, 70637-41. 

Sample 11. BRITISH HONDURAS: 
Cayo: Central Farm, MCZ 71608-9; El Cayo, 
UMMZ 75012; 1.6 km NW El Cayo, USNM 
71372-73; Xunantimich Ruins, MCZ 71619-21. 

Sample 12. GUATEMALA: El Peten: 



Tikal, UMMZ 117875(8), 117876(7); Uaxac- 
tim, AMNH 68507-16, 70938-42, UMMZ 
70415-19. 

Sample 13. GUATEMALA: El Peten: 
Piedras Negras, MCZ 66965, UIMNH 11363- 
93. 

Sample 14. GUATEMALA: Alta Verapaz: 
Canihor, UMMZ 91305(21), 91306(3), 91307 
(4), 91308; Finales, UMMZ 91316. 

Sample 15. GUATEMALA: Progreso: El 
Rancho, UMMZ 106994(11), 106995(6); 
Finca Bucural, UMMZ 107000(9), 107001(4). 

Sample 16. HONDURAS: Copdn: Go- 
pan, UMMZ 83035(16). 

Sample 17. HONDURAS: Atlantkla: 1 
km W La Ceiba, KU 101227; 2 km SE La 
Ceiba on Rio Cangrejal, KU 101228-38; 8 km 
SE La Ceiba on Rio Cangrejal, KU 101239-42; 
12 km SSE La Ceiba on Rio Cangrejal, KU 
101252; ca 15 km E La Ceiba in mountains 
above Corozal, JRM 2318, 2441-42, 2460-61, 
2470. 

Sample 18. HONDURAS: Yoro: 0.5 km 
N Coyoles, JRM 2004-7; 2 km S Coyoles 
along Rio Aguan, KU 101216-26, 101254-59, 
107911-14, 109973-74; 5 km E Coyoles, JRM 
2082-89; 25 km WSW Coyoles, Rancho San 
Lorenzo, JRM 2116. 

Sample 19. HONDURAS: Colon: 0.5 km 
SW Trujillo, JRM 2554; 1 km SSW Trujillo, 
KU 101249-51; 0.5-1.5 km W Trujillo, JRM 
2511; 1-3 km W Trujillo, KU 101243-47; 2 
km E Trujillo, jRM 2533-36; 2 km W Tru- 
jillo, KU 101248. 

Sample 20. HONDURAS: Olancho: 0.5- 
1.0 km WNW Catacamas, JRM 1433-37; 1 
km NW Catacamas, JRM 1551; 1.5 km NW 
Catacamas, JRM 1554-55; 2-3 km NW Cata- 
camas, JRM 1526-29; 4.5 km SE Catacamas, 
Esquela Agricola Nacional de Agricultura, 
JRM 1586-87, 1622, 1628-30; 6.5 km SE Cata- 
camas, JRM 1638-39. 

Sample 21. NICARAGUA: Zelat/a: Isla 
del Maiz Grande, AMNH 97045-67, 97637, 
MCZ 26970, 26972, 26974, USNM 94053, 
KU 85988, 85997, 85999, 101268, 101273-74, 
101276-77, 101281-86, 101292, 101299-301, 
101305-6, 101309-10. 

Sample 22. MfiXICO: Colima: Hacienda 
Paso del Rio, FMNH 75794, 126436, 126438- 
41, 126443-47, 126449-51, 167438, UMMZ 
80118(2), 80119; Manzanillo, FMNH 100054; 
1.6 km W Pascuales, UMMZ 80113(3); 0.4 
km E Pascuales, UMMZ 80114; 6 km SW 
Tecuman, UMMZ 80110; 8 km SW Tecuman, 
UMMZ 80111(3), 80112(5), 80120. 

Sample 23. MfiXICO: Colima: Colima, 
AMNH 5185, 15448-49; NW of Colima, 
AMNH 15838-42; 0.8 km SW Colima, AMNH 
12634-35; 2.4 km SW Colima, AMNH 12643; 
6.4 km SW Colima, KU 29529; near Salva- 
dor, MCZ 52172. 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



81 



Sample 24. MfiXICO: Guerrero: Mtns 
N of Acapulco, FMNH 126351, 126355, 
UIMNH 27344; 8 km E Coyuca, USNM 
133849-53; Between km 431-432, camp nr El 
Triente, FMNH 126353; 2 km N El Triente, 
UMMZ 119564; 1.6 km N Organos, S of El 
Triente, UIMNH 27345; E San Andreas de la 
Cruz, KU 87417. 

Sample 25. MEXICO: Guerrero: Aca- 
huizotla, TCWC 7780, 9587-88, 11320, 
UMMZ 119563; Agua del Obispo, UMMZ 
119565, TCWC 7575, KU 87413-16; nr Agua 
del Obispo between Rincon and Cajones, 
FMNH 126352, 126354, MCZ 56005; Mtns 
near Agua del OlMspo, TCWC 7576; 8 km SW 
Chilpancingo, TCWC 9590; 1.6 km SW Cot- 
otlipa, TCWC 9591. 

Sample 26. MEXICO: Oaxaca: 6.4 km S 
Candelaria, KU 38234; Chacalapa, KU 38230- 
33; 3.2 km W El Soledad, UIMNH 8421-27. 

San^ple 27. MEXICO: Oaxaca: Tres 
Cruces, 32 km SW Tehuantepec, UMMZ 
81897(2), 81898(3), 81899(2), 81900(3), 
81901(5), 81902(4), 81903(5), 81904(6). 

Sample 28. MEXICO: Oaxaca: 3.2 km 
E Tollocito, KU 39722-32, 39734, 44656-57. 

Sample 29. MEXICO: Oaxaca: Tapa- 
natepec, AMNH 80002-3, MCZ 27283-300; 
2.4 km S, 7.2 km E Tapanatepec, TCWC 
17047-48; 4 km W Tapanatepec, KU 59577- 
80; Rio Novillero, 4.3 km W Tapanatepec, 
UIMNH 39204-8. 

Sample 30. MEXICO: Chiapas: Finca 
San Bartola, ca 19 km SW Cintalapa, UIMNH 
8437-41, 39229-41; nr San Ricardo, FMNH 
106715, UIMNH 26157. 

Sample 31. MEXICO: Chiapas: Abel- 
lanal, 2 km SE Florida, TCWC 19764-65; 
Florida, 50 km E Altimirano, TCWC 19758- 
63, 19766-86; Las Tazas, 51 km E Altimirano, 
TCWC 19757. 

Sample 33. MEXICO: Chiapas: Esquint- 
la, LACM 9527-28; Distrito Soconusco, 6 km 
NE Esquintla, UMMZ 86862-71, 86873-88. 

Sample 34. MEXICO: Chiapas: Sabana 
de San Quintin, KU 94109-17, 94119. 

Sample 35. GUATEMALA: Solold: Fin- 
ca Santa Buenaventura, 1 km NW Panajachel, 
UMMZ 98215(2); Panajachel, MCZ 27515-20, 
KU 59581, 95653, UMMZ 98201(3), 98214 
(3); 1 km E Panajachel, KU 95657; 3 km E 
Panajachel, KU 95654-56; 3 km NE Pana- 
jachel, UMMZ 98202; 1 km NW Panajachel, 
UMMZ 98203(2), 98204; 2 km NE Pana- 
jachel, UMMZ 98212; 4 km NE Panajachel, 
UMMZ 98213; Solola, UMMZ 98205(2); On 
road to Solola, 1 km from Panajachel, UMMZ 
98206. 

Sample 36. GUATEMALA: Jutiapa: Fin- 
ca La Trinidad, UMMZ 107421, 107422(8), 
107423(5), 107424(3), 107425(6), 107427 
(3), 107428, 107429(3). 

Sample 37. GUATEMALA: Jutiapa: Ha- 



cienda Mongoy, UMMZ 107002(6), 107003, 
107004(3); jutiapa, UMMZ 106997(4), 
106998, 106999(16). 

Sample 38. EL SALVADOR: Libertad: 
2 km SE Colon, KU 62608-70; Laguna de 
Chanmico, UMMZ 117486(2); 0.8 km N, 16 
km W La Libertad, TCWC 17059; Rancho 
Belmar, nr La Libertad, LACM 9370. San 
Salvador: Instituto Tropical, San Salvador, 
KU 62066-67; 0.4 km NW Instituto Tropical, 
UMMZ 117487; 0.8 km NW Instituto Tropical, 
UMMZ 117488; 1.6 km NW San Salvador, 
KU 42262. 

Sample 39. NICARAGUA: Chinamle^a: 
4 km N, 2 km W Chichigalpa, KU 86036; 
Chinandega, MCZ 9540, 9546; Finca San 
Isidro, 10 km S Chinandega, KU 86030-31; 
Foothills, N Slope Volcan San Cristobal, KU 
86032-35; Hacienda Bellavista, Volcan Casita, 
KU 101875-77; San Antonio, KU 86008-26. 

Sample 40. NICARAGUA: Matagalpa: 
Guasquali, UMMZ 116420(3); Matagalpa, 
UMMZ 116414, 116418(5); 0.8 km E Mata- 
galpa, UMMZ 116416; 1.6 km W Matagalpa, 
UMMZ 116417, 116419; 2.4 km E Matagalpa, 
UMMZ 116415(2). 

Sample 41. COSTA RICA: Guanacasie: 
Hacienda Coyolar, 4.8 km N, 4.0 km W Li- 
beria, USC(CRE) 8206(15), 8207(2); Haci- 
enda La Norma, 5 km N, 4.5 km W Liberia 
on Rio Colorado, USC(CRE) 100(2), 103 
(3), 104, 105(3), 106(8), 107(4), 111, 251. 

Sample 42. COSTA RICA: Guanacaste: 
Canas, USNM 80894; 1.5 km E Canas, USC 
(CRE) 208-9; 1 km S Canas, USC(CRE) 
201; 7 km N, 3 km E Cai^as, USC (CRE) 270- 
77; Finca Taboga, 11 km S Caiias, USC 
(CRE) 631, 7166(2); Rio Javillo, 2 km S 
Las Cafias, USC (CRE), 202-3. 

Sample 43. COSTA RICA: Guanacaste: 
On first river N Santa Rosa, KU 40531-36; 
Road from Silencio to Tilaran, 1.8 km from 
Tilariin, USC (CRE) 6240; Road from Silen- 
cio to Tilaran, 2.6 km from Tilaran, USC 
(CRE) 6239(2); Tilaran, ANSP 24566-67; 
Rio Santa Rosa drainage 1.2 km by road SW 
Tilaran, USC (CRE) 7161; 1.6 km SW Tila- 
ran, USC(CRE) 6270; 2 km W Tilaran, USC 
(CRE) 732; 4.3 km by road NNE Tilaran, 
USC(CRE) 8082; 4 km ESE Los Angeles de 
Tilaran, KU 40527-28; 5.8 km from Tilaran 
on road to Los Angeles, USC (CRE) 6267; 
Troncadero, Laguna de Arenal, KU 40529-30. 

Sample 44. HONDURAS: Francisco- 
Morazdn: El Zamorano, Esquela Agricola 
Panamericana, AMNH 70368, MCZ 49765-67, 
10001-3, KU 101260-67; Rio Yeguare, AMNH 
70369-79; Valley of Rio Yeguare S of Teguci- 
galpa, AMNH 70470-73. 

Sample 46. MEXICO: Quintana Roo: 
Isla Mujeres, USNM 47568-70, 47651, KU 
70585-88. 



82 



MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



Sample 48. MEXICO: Michoacdn: Apat- 
zingan, FMNH 38977-85. 

Additional Specimens Not Used In STP 
Analysis. BRITISH HONDURAS: Belize: 
Aml:!ergris Cay, 3.2 km N San Pedro, MCZ 
71611; Belize, USNM 51879-80, 58373-78; 
Caijo: Augustine, Mountain Pine Ridge, MCZ 
71622-24; Corozal: 0.3 km N Corozal, MCZ 
71610; Orange Walk: Gallon Jug, MCZ 
71612-18; Stann Creek: Mango Creek, MCZ 
71627-28; 8 km SW Mango Creek, MCZ 
71626; 6.4 km S Waha Leaf Creek, MCZ 
71625; Toledo: 1.6 km W Svvazey Bridge, 
Monkey River, MCZ 71629. COSTA RICA: 
Alajuela: San Mateo, USNM 37491; "Auto- 
pista-AJajuela," UCR uncatalogued; Cartage: 
La Carpintera, AMNH 16314, 16316; Guana- 
caste: Bebedero, USC(CRE) 7162; 1.6 km 
N Guayabo de Bagaces, USC(CRE) 7024; 
Playa Samara, outer coast of Nicoya Peninsula, 
USC(CRE) 8252; Parrita, La Julieta, Finca 
La Ligia, USC(CRE) 8255(4), 8260, 8262; 
1.6 km S Santa Cruz, USC(CRE) 8218; 1.8 
km N Santa Cruz, USC(CRE) 8217(2); 
Puntarenas: 4.3 km E Esparta, UCR 547; San 
Jose: Barrio Mexico, San Jose, KU 40526. 
EL SALVADOR: La Paz: 3 km E San Ra- 
fael Obrajuela, KU 62065; San Miguel: San 
Pedro, MCZ 57081. GUATEMALA: Guate- 
mala: El Rosario, FMNH 68708-10; Izabal: 
1.6 km NE York, KU 59574-75; San Marcos: 
Finca Carolina, MCZ 22144-48, 27301-5; San- 
to Rosa: Finca Las Vinas, UMMZ 107073-74; 
Departamento Unknotvn: MCZ 5831, 99840- 
41. HONDURAS: Colon: Belfate, AMNH 
58607-9; Comaijagua: La Libertad, MCZ 
38923; Copdn: Hugito, ANSP 22195-98; Cor- 
tes: E side Lago de Yojoa, KU 67332; Rancho 
Agua Azul, Lago de Yojoa, MCZ 49964; 3.2 
km W San Pedro Sula, MCZ 29387-88; El 
Paraiso: Arenal, 25 km E Jalapa ( Nicaragua ) , 
UCLA 14759; Valle de Jamastran, AMNH 
70333-35; Francisco-Morazdn: Cantarranas. 
ANSP 22199-212; Gracias a Dios: Tancin, ca 
15 km NW Puerto Lempira, JRM 1732, 1752. 
MEXICO: Campeche: 3.2 km E Barro San 
Pedro Y Pablo, AMNH 88877; Champoton, 
UMMZ 72956-57; Chiapas: 3.2 km E El Real, 



KU 43663-64; Palenque, MCZ 66966; Across 
Rio Usimiacinta from Piedras Negras (Guate- 
mala), USNM 108600; San Jnanita, Palenque, 
USNM 108601; Colima: Queseria, UMMZ 
80109; nr Queseria, FMNH 126437, 126442, 
126448; Guerrero: nr Rincon, FMNH 100080; 
Oaxaca: La Concepcion, nr Tehuantepec, 
AMNH 66932; Cosolapa, USNM 133920; Ju- 
quila Mixes, AMNH 89761-65; Mitla, AMNH 
91023-24; 9.6 km N Putla, UIMNH .52952-53; 
Rio Cacahuatepec, UIMNH 529,50; Rio Canoa, 
26 km W Pinotepec Nacional, UIMNH .52951; 
Sta. Lucia, Tehuantepec, AMNH 65077-78, 
65085; Tehuantepec, USNM 30163; Zacatepec, 
UIMNH 52948-49; Puehla: Necapa, Rio Ne- 
capa, AMNH 76441-43; nr Vegas de Xuchil, 
AMNH 58217; Quintana Roo: 8 km inland 
from Vigia, Ascension Bay, UMMZ 78586; 
San Luis Potosi: Ciudad Maiz, FMNH 126942; 
nr Cuidad Maiz, FMNH 126946; Palictla, 
AMNH 67358; 8 km S Valles, FMNH 126947; 
Tabasco: La Venta, USNM 117350; Tamau- 
lipas-: S of Antiguo Morelos, FMNH 126944- 
45; Hacienda La Clementina, FMNH 126943, 
126948, MCZ 56006, USNM 106141-42; 
Rancho Santa Ana, 12.8 km E Padilla, MCZ 
43634; Rio Guayalejo, nr Magiscatzin, MCZ 
45557, UMMZ 88232; 11 km W Victoria, 
FMNH 100050; Veracruz: Jalapa, FMNH 
126706; Orizaba, USNM 133839; Panuco, 
MCZ 19255-59; 4 km E Papantla, KU 24189, 
24194; San Andres Tuxtla, FMNH 100020; 16 
km WNW Temapache, KU 61791-92; Yuca- 
tan: Chichen Itza, MCZ 52171; Culebra Cay, 
Ascencion Bay, UMMZ 78587; Progreso, 
FMNH 100030; Estado Unktwwn: "Mexico," 
( Restricted to Tehuantepec, Oaxaca, by Smith, 
1940), ZMB 868. NICARAGUA: Esteli: 
Finca Daraili, 5 km N, 14 km E Condega, KU 
85984; Madriz: 1.6 km SE Yalaguina, TCWC 
17051; Managua: 12 km E Managua, UCLA 
14787; 6.4 km E San Lorenzo, KU 42249; 
Rio San Juan: Tuli Creek, AMNH 16675; 
Rivas: within 3 km of Moyogalpa, Isla Ome- 
tepe, KU 86029; Zelaija: Huaunta Haulover, 
ANSP 15438-41; Departamento Unknown: 
ANSP 9133-34. 



ECHTERNACHT: MIDDLE AMERICAN LIZARDS 



83 



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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 



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The manuscript for this publication was 
completed in March, 1970, and submitted as a 
dissertation in partial fulfillment of the re- 
quirements for the PhD at the University of 
Kansas. 




3 2044 093 361 616 



Date Due 




BOUND 



1972